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Turczaninowia 2008, 11(1) : 5–168

УДК 582.594 (597)

L. Averyanov Л.В. Аверьянов

THE ORCHIDS OF VIETNAM

ILLUSTRATED SURVEY

Part 1

SUBFAMILIES APOSTASIOIDEAE, CYPRIPEDIOIDEAE

AND SPIRANTHOIDEAE

The article opens serial publication of illustrated critical taxonomical survey of

orchids in the flora of Vietnam. The first part of this monograph includes introduction,

illustrated glossary of terminology used in identification keys and in description of

taxa, key for identification of genera, as well as taxonomical treatment of three

subfamilies – Apostasioideae, Cypripedioideae and Spiranthoideae with 21 genera

and 85 species. Valid name, necessary synonyms, type material citation, short

description, data on ecology and distribution, as well as list of studied voucher specimens

for each species are provided. Mentioned species and varieties are illustrated with

line drawings and color photographs.

PREFACE

Basic primary knowledge on orchid taxonomy and diversity in Eastern Indochina

and in Vietnam was obtained from landmark surveys (Gagnepain, Guillaumin, 1943;

Pham-hoang, 1960, 1972, 1993, 2000; Averyanov, 1990, 1994; Seidenfaden, 1992) based

mainly on collections of “French” and post-war periods housed now in largest part in

Herbaria of Paris (P) and Saint Petersburg (LE). Mentioned publications were used

in some compilative reviews few later (Tran Hop, 1998; Nguyen Thien Tich, 2001;

Nguyen Tien Ban, Averyanov, Duong Duc Huen, 2005). According to most recent

assessment (Averyanov, Averyanova, 2003) flora of Vietnam includes about 158 genera

and 900 discovered orchid species, which may be estimated as about 80% of all orchids

occurring on the territory of the country. Meanwhile, recent exploration of numerous

earlier inaccessible territories of the country brings last time many more new

discoveries, which essentially expand our knowledge about regional plant diversity.

Illustrated taxonomical survey of all species hitherto reported for Vietnam including

most recent discoveries based on collected voucher herbarium specimens is the main

goal of presented monograph.

Monograph treatment will successively follow family system of R. Dressler

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

(Dressler, 1993) with few original additions and amendments. The orchid family system

in used form is presented below as follow:

ORCHIDACEAE Juss.

Subfam. 1. Apostasioideae Reichenb.

Apostasia Blume, Neuwiedia Blume

Subfam. 2. Cypripedioideae Lindl.

Paphiopedilum Pfitz.

Subfam. 3. Spiranthoideae Dressler

Trib. 3.1. Tropidieae Dressler

Corymborkis Thouars, Tropidia Lindl.

Trib. 3.2. Cranichideae Endlicher

Subtrib. 3.2.1. Goodyerinae Klotzsch

Anoectochilus Blume, Cheirostylis Blume, Erythrodes Blume, Goodyera

R. Br., Herpysma Lindl., Hetaeria Blume, Ludisia A. Rich., Macodes Lindl.,

Myrmechis Blume, Odontochilus Blume, Orchipedum Breda, Rhomboda Lindl.,

Vrydagzynea Blume, Zeuxine Lindl., Zeuxinella Aver.

Subtrib. 3.2.2. Spiranthinae Lindl.

Spiranthes L.C. Rich

Subfam. 4. Orchidoideae

Trib. 4.1. Diurideae Endlicher

Subtrib. 4.1.1. Acianthinae Schlecht.

Corybas Salisb.

Subtrib. 4.1.2. Cryptostylidinae Schlecht.

Cryptostylis R. Br.

Trib. 4.2. Orchideae

Subtrib. 4.2.1. Orchidinae

Amitostigma Schlecht., Brachycorythis Lindl., Hemipilia Lindl., Platanthera

L.C. Rich., Vietorchis Aver. et Averyanova

Subtrib. 4.2.2. Habenariinae Benth.

Diplomeris D. Don, Habenaria Willd., Herminium R. Br., Pecteilis Rafin.,

Peristylus Blume

Subfam. 5. Epidendroideae Lindl.

Group of primitive tribes

Trib. 5.1. Neottieae Lindl.

Aphyllorchis Blume, Epipactis Sw., Listera R. Br.

Trib. 5.2. Vanilleae Blume

Subtrib. 5.2.1. Galeolinae Garay

Cyrtosia Blume, Erythrorchis Blume, Galeola Lour.


Turczaninowia 2008, 11(1) : 5–168

Subtrib. 5.2.2. Vanillinae Lindl.

Vanilla Sw.

Subtrib. 5.2.3. Lecanorchidinae Dressler

Lecanorchis Blume

Trib. 5.3. Gastrodieae Lindl.

Subtrib. 5.3.1. Gastrodiinae Lindl.

Didymoplexiella Garay, Didymoplexiopsis Seidenf., Didymoplexis Griff.,

Gastrodia R. Br., Yoania Maxim.

Subtrib. 5.3.2. Epipogiinae Schlecht.

Epipogium R. Br., Stereosandra Blume

Trib. 5.4. Nervilieae Dressler

Nervilia Gaudich.

Epidendroid group

Trib. 5.5. Arethuseae Lindl.

Subtrib. 5.5.1. Arundinae Dressler

Arundina Blume

Subtrib. 5.5.2. Bletiinae Benth.

Acanthephippium Blume, Anthogonium Lindl., Bletia Ruiz et Pavon, Calanthe

R. Br., Cephalantheropsis Guillaum., Eriodes Seidenf., Hancockia Rolfe,

Mischobulbon Schlecht., Nephelaphyllum Blume, Pachystoma Blume, Phaius Lour.,

Plocoglottis Blume, Spathoglottis Blume, Tainia Blume

Trib. 5.6. Malaxideae Lindl.

Subtrib. 5.6.1. Malaxidinae Benth. et Hook. f.

Liparis L.C. Rich., Malaxis Sw.

Subtrib. 5.6.2. Oberoniinae Aver.

Hippeophyllum Schlecht., Oberonia Lindl.

Trib. 5.7. Coelogyneae Pfitz.

Subtrib. 5.7.1. Thuniinae Schlecht.

Thunia Reichenb.

Subtrib. 5.7.2. Coelogyninae Bentham

Coelogyne Lindl., Neogyna Reichenb. f., Otochilus Lindl., Panisea Steud.,

Pholidota Hook., Pleione D. Don

Trib. 5.8. Epidendreae Humb., Bonpl. et Kunth

Subtrib. 5.8.1. Glomerinae Schlecht.

Agrostophyllum Blume

Subtrib. 5.8.2. Polystachyinae Pfitz.

Polystachya Hook.

Dendrobioid group

Trib. 5.9. Podochileae Pfitz.

Subtrib. 5.9.1. Eriinae Benth.

Callostylis Blume, Ceratostylis Blume, Cryptochilus Wall., Eria Lindl.,

Porpax Lindl., Trichotosia Blume

Subtrib. 5.9.2. Podochilinae Benth. et Hook.

Appendicula Blume, Podochilus Blume

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Subtrib. 5.9.3. Thelasiinae Schlecht.

Phreatia Lindl., Thelasis Blume

Subtrib. 5.9.4. Dendrobiinae Lindl.

Dendrobium Sw., Epigeneium Gagnep., Flickingeria Hawkes

Subtrib. 5.9.5. Bulbophyllinae Schlecht.

Bulbophyllum Thouars, Hamularia Aver., Monomeria Lindl., Sunipia Smith,

Trias Lindl.

Cymbidioid group

Trib. 5.10. Cymbidieae Pfitz.

Subtrib. 5.10.1. Bromheadiinae Dressler

Bromheadia Lindl.

Subtrib. 5.10.2. Eulophiinae Benth.

Eulophia R. Br., Dipodium R. Br., Geodorum Jackson

Subtrib. 5.10.3. Collabiinae Schlecht.

Chrysoglossum Blume, Collabium Blume, Diglyphosa Blume

Subtrib. 5.10.4. Thecostelinae Schlecht.

Thecopus Seidenf., Thecostele Reichenb.

Subtrib. 5.10.5. Cymbidiinae Benth.

Cremastra Lindl., Cymbidium Sw.

Subtrib. 5.10.6. Acriopsidinae Dressler

Acriopsis Blume

Subfam. 6. Vandoideae

Trib. 6.1. Vandeae Lindl.

Subtrib. 6.1.1. Aeridinae Pfitz.

Acampe Lindl., Adenoncos Blume, Aerides Lour., Arachnis Blume,

Ascocentrum Schlecht., Ascocentropsis Senghas & Schildh., Biermannia King et Pantl.,

Brachypeza Garay, Chiloschista Lindl., Christensonia Haager, Cleisocentron Bruehl,

Cleisomeria G. Don, Cleisostoma Blume, Cleisostomopsis Seidenf., Cryptopylos Garay,

Diploprora Hook. f., Doritis Lindl., Eparmatostigma Garay, Esmeralda Reichenb.,

Gastrochilus D. Don, Grossourdya Reichenb., Holcoglossum Schlecht., Hygrochilus

Pfitz., Kingidium P.F. Hunt, Luisia Gaudich., Malleola J.J. Smith, Micropera Lindl.,

Microsaccus Blume, Ornithochilus Lindl., Papilionanthe Schlecht., Parapteroceras

Aver., Pelatantheria Ridl., Pennilabium J.J. Smith, Phalaenopsis Blume, Pomatocalpa

Breda, Pteroceras Hassk., Renanthera Lour., Rhynchostylis Blume, Rhynchogyna

Seidenf. et Garay, Robiquetia Gaudich., Saccolabiopsis J.J. Smith, Sarcoglyphis Garay,

Schoenorchis Blume, Smitinandia Holttum, Stereochilus Lindl., Staurochilus Pfitz.,

Taeniophyllum Blume, Thrixspermum Lour., Trichoglottis Blume, Tuberolabium

Yamamoto, Uncifera Lindl., Vanda Jones, Vandopsis Pfitz.

Illustrated survey of three subfamilies – Apostasioideae, Cypripedioideae

and Spiranthoideae is presented here in form of standard taxonomic treatment, which

includes identification keys for all mentioned taxa and their short characterization.

Correct name (with standard taxonomic reference), type, data about volume and

distribution is reported for each taxonomic group. Data for each genus also include


Turczaninowia 2008, 11(1) : 5–168

short description, total number of species and number of species in the flora of Vietnam

(figures in brackets), as well as the genus distribution.

For each species (subspecies or variety) the following information is reported:

– valid name, most significant synonyms and citation of most important recent

monographs,

– all available data about type,

– short description,

– available data on ecology, elevation of observed habitats, phenology, frequency

of occurrence in the nature with approximate estimation of species status

according to IUCN categories for the territory of Vietnam,

– general distribution and distribution in Vietnam (provinces where species

was reported from are mentioned in alphabetical order in brackets ),

– list of studied verified specimens and index of their host herbaria,

– when necessary short notices on species taxonomy, biology, ecology,

phenology or variation are also provided.

– line drawings or (and) color photographs accompany in the book each species

reported for the flora of Vietnam; collecting numbers of plants used as a model for

illustration are cited on drawings or in notices to photographs.

Text of labels is extremely abbreviated whenever being cited. It usually includes

only province name, district name, collectors name and collector number, or, if necessary

date of collection. Largest recent collections are abbreviated and designated as series

with following prefixes:

CBL – Cao Bang Limestone – collections on program of U.S.A. National

Geographic Society “Limestone Flora of Cao Bang Province of northern Vietnam”

(years 1998-1999, # 6300-98) with principal investigator Dr. Nguyen Tien Hiep,

CPNP – collections in Cuc Phuong national park (commonly without indication

of collectors),

DDS – Prof. D.D. Soejarto – collections according to International Cooperative

Biodiversity Groups program with this person as principal investigator,

DKH – Dr. D. Harder – collections according to expeditions with this person

as a principal investigator,

HAL – Dr. Nguyen Tien Hiep, Prof. Leonid V. Averyanov, Prof. Phan Ke

Loc – collections in collaborative explorations of these persons,

HLF – Henry Luce Foundation, collections of different collectors according

to Vietnam Botanical Conservation Program supported from Henry Luce Foundation,

LX-VN – collections of Soviet-Vietnamese Expedition (commonly without

exact indication of collectors),

NMC – collections of staff member of Cuc Phuong national park – Mr. Nguyen

Manh Cuong on the territory of the national park,

NTH – Dr. Nguyen Tien Hiep – collections in expeditions with this person as

principal investigator,

P – Prof. Phan Ke Loc – collections in expeditions with this person as principal

investigator,

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

VA - Vietnam-American Series – collections of international group with Dr.

N.T. Hiep as principal investigator.

VH – Vietnamese Highlands – collections on program of U.S.A. National

Geographic Society “Flora of Highlands of South Vietnam” (years 1993-2001, # 5094-

93, 5803-96, 6383-98) with principal investigator Prof. Leonid V. Averyanov.

WP – Vietnam-Chinese expedition of 2003 year with Prof. Phan Ke Loc as

co-leader from Vietnamese Side.

Modern administrative divisions of Vietnam into provinces and city territories,

as well as list of their names, which are used in indication of species distribution within

Vietnam, are presented on map 1.

Illustrated glossary of orchid terminology used in identification and plant

descriptions are presented below in the next chapter.

ILLUSTRATED GLOSSARY OF THE ORCHID

IDENTIFICATION TERMINOLOGY

Additional explanations of terms are available on cited figures (Fig. 1–12).

Abaxial – the side away from the axis, normally the lower surface.

Acaulescent – becoming stemless.

Names of provinces, names of city areas and figures designating these units on the map:

1 – Dien Bien

2 – Lai Chau

3 – Lao Cai

4 – Yen Bai

5 – Son La

6 – Ha Giang

7 – Cao Bang

8 – Tuyen Quang

9 – Bac Can

10 – Phu Tho

11 – Thai Nguyen

12 – Lang Son

13 – Bac Giang

14 – Quang Ninh

15 – Vinh Phuc

16 – Ha Noi city

17 – Bac Ninh

18 – Ha Tay

19 – Hung Yen

20 – Hai Duong

21 – Hai Phong

22 – Hoa Binh

23 – Ha Nam

24 – Thai Binh

25 – Ninh Binh

26 – Nam Dinh

27 – Thanh Hoa

28 – Nghe An

29 – Ha Tinh

30 – Quang Binh

31 – Quang Tri

32 – Thua Thien-Hue

33 – Da Nang city

34 – Quang Nam

35 – Kon Tum

36 – Quang Ngai

37 – Gia Lai

38 – Binh Dinh

39 – Dak Lak

40 – Phu Yen

41 – Dak Nong

42 – Khanh Hoa

43 – Binh Phuoc

44 – Lam Dong

45 – Ninh Thuan

46 – Tay Ninh

47 – Binh Duong

48 – Dong Nai

49 – Binh Thuan

50 – Long An

51 – Ho Chi Min city

52 – Ba Ria-Vung Tau

53 – An Giang

54 – Dong Thap

55 – Tien Giang

56 – Kien Giang

57 – Can Tho

58 – Vinh Long

59 – Ben Tre

60 – Hau Giang

61 – Tra Vinh

62 – Ca Mau

63 – Bac Lieu

64 – Soc Trang


Turczaninowia 2008, 11(1) : 5–168

Map 1. Modern administrative division of Vietnam territory (for 2007 year) (except Hoang Sa

and Truong Sa Archipelagos).

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Acaulous – stemless or nearly so.

Achlorophyllous – without chlorophyll, as in plants or plant structures lacking

green color.

Actinomorphic – radially symmetrical.

Aculeate – prickly; covered with prickles.

Acuminate – having a gradually tapering point. Fig. 12.

Acute – distinctly and sharply pointed, but not drawn out. Fig. 12.

Adaxial – the side toward the axis, normally the upper surface.

Adnate – united or joined together.

Adpressed (appressed) – lying flat for the whole length of the organ.

Amplexicaul – clasping the stem.

Androclinium – see clinandrium.

Androecium – all of the stamens in a flower.

Anther – the expanded, apical, pollen bearing portion of the stamen. Fig. 4–7, 10.

Anther cup (operculum) – highly modified, easily removable lid-like anther

wall, which covers pollinia placed on the top of column in many representatives of

Dendrobioideae and Vandoideae subfamilies. Fig. 8–10.

Antrorse – directed forward or upward.

Aphyllous – without leaves.

Apiculate – with a short, sharp, but not stiff, point. Fig. 12.

Appressed – see adpressed.

Approximate – drawn close together, but not united.

Aristate – awned, sharply terminated by a hard, straight point. Fig. 12.

Articulate – jointed. Fig. 3.

Articulation – point of attachment. Fig. 3.

Attenuate – tapering gradually to a narrow base. Fig. 12.

Auricle 1 – a small lateral outgrowth on the top of column in representatives

of subfamily Orchidoideae. Fig. 7.

Auricle 2 – a small, ear-shaped appendage, commonly at the base of lip or

leaf. Fig. 12.

Auriculate – with a small ear-like lobe. Fig. 12.

Axial placentation – placentation when ovules are attached to the central

axis of an ovary with two or more locules. Fig. 2, a.

Axil – the point at the angle between a leaf and a stem.

Axillary – borne in the axil.

Bark epiphytes (trunk epiphytes) – epiphytes growing on branches of middle

size in medium zone of tree canopies. Fig. 1, f.

Berry – a fleshy, many-seeded fruit composed of more than one carpel.

Bicuspidate – having 2 sharp points.

Bidentate – with two teeth. Fig. 12.

Bifid – divided into 2 shallow segments, usually at the apex. Fig. 12.

Bifurcate – forked.

Bipartite – divided nearly to the base into two portions.


Turczaninowia 2008, 11(1) : 5–168

Fig. 1. Ecological groups of orchids and their habitats.

Orchids in the flora of Vietnam may be terrestrial plants (a, about 20%), achlorophyllous

obligate mycotrophic (saprophytic) herbs (b, about 2%), lithophytes (c, about 8%), lianas or

vines (d, about 2%), humus epiphytes (e, about 15%), bark epiphytes (f, about 48%) and

branch epiphytes (g, about 5%).

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 2. Basic orchid morphology: a, b – transversal section of ovary with axial placentation in

Apostasia odorata (a) and parietal placentation in Dendrobium cariniferum (b); c-e – main

types of orchid growth system with monopodial shoot (c), sympodial shoot with apical inflorescence

(d) and sympodial shoot with lateral inflorescence (e); rings designate flowers, filled

rings designate fruits, lines designate bracts/leaves, arrows show direction of growth, broken

line designate obsolete old part of shoot; f-i – main types of orchid inflorescences according

to features of their development: heteranthous in Coelogyne cristata (f), proteranthous in

C. lentiginosa (g), synanthous in Pholidota rubra (h) and hysteranthous (i) inflorescence in

C. fimbriata; j, k – types of leaf development (transversal section of developing leaves) with

convolute leaves in Liparis nervosa (j) and duplicate leaves in C. fimbriata (k); l-n – types of

orchid leaves (transversal section of mature leaves) with plicate in Tainia viridifusca (l),

conduplicate in Paphiopedilum concolor (m) and laterally flattened leaves in Oberonia

ensiformis (n).


Turczaninowia 2008, 11(1) : 5–168

Bract – more or less reduced leaf-like organ bearing a flower, inflorescence

or partial inflorescence in its axil. Fig. 3.

Branch epiphytes (canopy epiphytes) – epiphytes growing on small branchlets

and perennial leaves in peripheral zone of tree canopies. Fig. 1, g.

Canopy epiphytes – branch epiphytes.

Broadly elliptic – spindle-shaped, two-dimensional; with an index from 1 to

about 1.4. Fig. 11.

Broadly lanceolate – lance-shaped, narrow, tapering toward the apex more

than toward the base; with an index from 3 to around 4. Fig. 11.

Broadly oblanceolate – inversely broadly lanceolate. Fig. 11.

Broadly obovate – inversely broadly ovate. Fig. 11.

Broadly ovate – egg-shaped in outline, broader at the base, two-dimensional;

with an index less than 1.4. Fig. 11.

Broadly triangular – three-angled, with an index less than 0.85. Fig. 11.

Bursicle – a cup-like covering over the viscidium in some representatives of

subtribe Orchidinae. Fig. 10.

Caducous – falling off early, like floral bracts in some species of Calanthe or

Coelogyne.

Caespitose – tufted.

Callus (pl. calli) – a thickening or protuberance, commonly on the lip. Fig. 6.

Calyculus – a small cup or circle of bract-like structures outside of the sepals

in species of Lecanorchis. Fig. 8.

Campanulate – bell-shaped.

Canaliculate – channeled, with a longitudinal groove.

Canopy epiphytes – see branch epiphytes. Fig. 1, g.

Capilletium – the sterile kapok-like hairs within the fruit mellowing seed

masses, particularly in epiphytes. Fig. 9.

Capitate – head-like.

Capsule – a dry, many-seeded, dehiscent fruit composed of more than one

carpel. Fig. 9.

Carinate – keeled.

Carpel – highly modified megasporophyll, formed from one modified leaf

bearing ovules (megasporangia) and seeds; in the orchids, the three carpels are so

united and modified as to be nearly unrecognizable. Fig. 2, a, b.

Caudate – tailed; with a tail-like appendage. Fig. 12.

Caudicle – a slender elastic extension of the pollinium, or a mealy portion at one

end of the pollinium; the structure is a part of the pollen, which is formed within the

anther. Fig. 6, 7, 10.

Central – relating to the central part of a body.

Ciliate – having fine hairs at the margin, refers usually to the margin of leaf or

tepal. Fig. 12.

Clavate – club-shaped, pointed at one end and rounded at the other.

Claw – the conspicuously narrowed and attenuate base of an organ.

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Fig. 3. Structure of orchid shoot.

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.


Turczaninowia 2008, 11(1) : 5–168

Fig. 4. Orchid morphology (subfamily Apostasioideae).

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Cleistogamous – refers to flowers which regularly self-pollinate without

opening.

Clinandrium (androclinium) – the anther bed, that part of the column under,

or surrounding, the anther. Fig. 9.

Column – more or less modified structure formed by union of filaments and

style. Fig. 4–6, 8–10.

Column wing – a wing- or arm-like appendage of the column, usually lateral.

Columnfoot – a ventral extension of the base of the column which has the lip

attached at its tip. Fig. 8.

Complanate – flattened or compressed.

Conduplicate – leaf or leaf-like organs, with a single median fold, with each

half being flat. Fig. 2, m.

Congeneric – belong to the same genus.

Connate – united.

Connective – the sterile portion of the filament connecting the two pollen

sacs of an anther. Fig. 7, 10.

Connivent – coming into close contact or converging, but not united.

Conspecific – belonging to the same species.

Convolute – rolled up longitudinally with parts in an overlapping arrangement.

Fig. 2, j.

Cordate – heart-shaped with the base at the broad, notched end. Fig. 11, 12.

Coriaceous – leathery.

Corolla – the inner whorl of the perianth, petals.

Corymb – flat-topped inflorescence.

Crenate – scalloped, toothed with crenations. Fig. 11.

Crenulate – crenate, but the teeth small. Fig. 11.

Cristate – crested.

Cucullate – hooded, hood-shaped.

Cuneate – wedge-shaped. Fig. 12.

Cuspidate – tipped with a sharp, rigid point. Fig. 12.

Decurved – curved or bent down.

Deflexed – bent outwards.

Dentate – toothed. Fig. 11.

Denticulate – minutely toothed. Fig. 11.

Determinate – a habit of growth in which each unit has a limited growth,

usually in sympodial shoot.

Disc – the face of any flat organ with special reference to the lip.

Distal – away from the base, toward the apex.

Distichous – having leaves or other organs in two opposite rows.

Dorsal – refers to the back or outward surface of an organ in relation to the

axis; in orchids refers to the upper side of the flower; see abaxial.

Dorsal sepal – see median sepal.

Dorsiventral – an organ which has more or less distinct dorsal and ventral


Turczaninowia 2008, 11(1) : 5–168

Fig. 5. Orchid morphology (subfamily Cypripedioideae).

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Fig. 6. Orchid morphology (subfamily Spiranthoideae).

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.


Turczaninowia 2008, 11(1) : 5–168

surfaces, as a non-terete leaf.

Duplicate – refers to the folding of leaves during development, folded once

with each half flat. Fig. 2, k.

Echinate – with numerous stiff hairs or spines.

Elastoviscin – a very elastic substance found in pollinia and especially in

caudicles.

Ellipsoid – spindle-shaped; narrow and tapering at the ends, three-dimensional.

Elliptic – spindle-shaped, two-dimensional; with an index around 1.4–3. Fig. 11.

Emarginate – notched, usually at the apex. Fig. 12.

Endemic – confined to a given region.

Ensiform – sword-shaped. Fig. 11.

Entire – simple and with a smooth edge, as contrasted to toothed or lobed.

Fig. 11.

Ephemer – living a very short time (usually annual plants of arid areas).

Ephemeral – of flowers, referring to those which last only a few hours.

Ephemeroid – perennial plants of arid areas with very short vegetative and

reproductive periods.

Epichile – the terminal part of a complex lip when it is distant from the basal

portion. Fig. 6.

Epigeous – growing upon the ground.

Epiphyte – a plant growing on another plant but not parasitic. Fig. 1, e-g.

Equitant – said of conduplicate or laterally flattened leaves or bracts which

overlap each other in two ranks.

Erose – bitten or gnawed. Fig. 11.

Extrafloral – occurring outside of a flower; refers usually to glands that are

found outside of flower buds.

Falcate – sickle-shaped. Fig. 11.

Farinose – mealy.

Fasciculate – clustered or bundled.

Filament – the stalk of the stamen which bears the anther. Fig. 4, 5.

Filiform – thread-like.

Fimbriate – fringed, refers usually to the margin of leaf or tepal. Fig. 11.

Flabellate – fan-shaped. Fig. 11.

Flexuose – bent alternately in opposite directions.

Flora – the array of all plant species occurring in a region.

Floral bract – more or less reduced leaf-like organ bearing a flower (or

reduced floral bud) in its axil. Fig. 3, 5, 6, 8.

Fovea – a small depression or pit, like in lip of Malaxis species.

Fusiform – spindle-shaped.

Geniculate – abruptly bent like a knee-joint.

Geophyte – plants, in which new annual shoot develops from underground

bud, usually tuberiferous or bulb ephemeroids.

Gibbous – swollen or enlarged on one side; ventricose.

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Fig. 7. Orchid morphology (subfamily Orchidoideae).

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.


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Fig. 8. Orchid morphology (subfamily Epidendroideae).

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Glabrous – hairless.

Gland 1 – an appendage, protuberance, or other structure which secretes

nectary substances. Fig. 6.

Gland 2 – see viscidium.

Globose – spherical.

Gynostemium – see column.

Hamulus – a pollinium stalk which is formed by modified bended rostellum

apex. Fig. 10.

Hastate – spear-shaped, with the basal lobes turned outwards. Fig. 11, 12.

Heteranthous – an apical inflorescence produced on a separate shoot which

does not develop to produce a pseudobulb and leaves. Fig. 2, f.

Hirsute – hairy.

Hispid – bristly.

Humus epiphytes – epiphytes growing in forks of tree stem on thick

accumulations of humus. Fig. 1, e.

Hyaline – glass-like, colorless, translucent.

Hygroscopic – capable of absorbing moisture from the atmosphere.

Hypochile – the basal portion of a complex lip. Fig. 6, 9.

Hysteranthous – an apical inflorescence produced after the pseudobulbs

and leaves have developed. Fig. 2, i.

Imbricate – overlapping.

Incumbent – lying on or against.

Indehiscent – not opening at maturity along definite lines or by pores.

Index (leaf, tepal index) – the length of flat figure divided by its width.

Indumentum – any covering, commonly refers to hairiness.

Inferior – inserted below, like the ovary in the orchids.

Inflorescence – the flowering part of a plant, which includes usually floral

axis, floral bracts and flowers. Fig. 3, 5–9.

Inflorescence bract – more or less reduced leaf-like organ bearing an

inflorescence or partial inflorescence (or reduced inflorescence bud) in its axil. Fig. 3, 7.

Internode – the section of a stem between two nodes. Fig. 3.

Isthmus – a narrow portion of a lip or petal.

Keel – a prominent longitudinal ridge, like the keel of a boat. Fig. 6, 8, 9.

Labellum – see lip.

Lacerate – cut or cleft irregularly, refers usually to the margin of leaf or

tepal. Fig. 11.

Lacinate – cut into narrow, irregular lobes or segments, refers usually to the

margin of leaf or tepal. Fig. 11.

Lamella – a membrane or septum.

Lamina – a blade, the expanded flat portion of a leaf or tepal.

Lanceolate – lance-shaped, narrow, tapering toward the apex more than

toward the base; with an index around 4–7. Fig. 11.

Lateral – borne on or near the side of midline of a bilaterally symmetrical organ.


Turczaninowia 2008, 11(1) : 5–168

Lateral sepals – adaxial segments of the outer whorl of perianth (calyx).

Fig. 6–9.

Laterally flattened – compressed from lateral sides.

Lax – loose, distant.

Leaf blade – the broad part of a leaf. Fig. 3.

Leaf index – see index.

Leaf sheath – the leaf base which surrounds the stem. Fig. 3, 5, 6.

Liana – a plant with long stem not self-supporting, but climbing or trailing on

any erect substratum. Fig. 1, d.

Ligulate – with a ligule; strap- or tongue-shaped.

Ligule – a strap-like body.

Linear – leaf blade at least 12 times longer than broad, with the sides parallel.

Fig. 11.

Lingulate – tongue-shaped.

Lip – in orchids highly modified median (adaxial) segment of the inner whorl

of perianth (corolla). Fig. 4–9.

Lithophyte – growing on stones and rocks. Fig. 1, c.

Lobule – a small lobe.

Longitudinal section (sagittal section) – section along axis of an organ.

Lunate – half-moon shaped.

Massula (pl. massulae) – a packet of pollen.

Median – situated on the midline of a bilaterally symmetrical organ.

Median sepal – abaxial segment of the outer whorl of perianth (calyx). Fig. 5–9.

Mentum – a chin-like projection formed by the lateral sepals and extended

columnfoot. Fig. 8.

Mesochile – the middle portion of a complex lip. Fig. 6.

Monopodial growth – indefinite growth which continues from a terminal

bud from season to season. Fig. 2, c.

Mucro – a sharp terminal point. Fig. 12.

Mucronate – tipped with a short, sharp, abrupt point (mucro). Fig. 12.

Mycorrhiza – a symbiotic relationship between a fungus and the root of a plant.

Mycotrophic (saprophytic) – plants which obtain nutrients from fungi by

means of mycorrhizal symbiotic or parasitic relationships. Fig. 1, b.

Myrmecophyte – any plant which is associated with ants.

Narrowly elliptic – spindle-shaped, two-dimensional; with an index around 3-

8. Fig. 11.

Narrowly lanceolate – lance-shaped, narrow, tapering toward the apex more

than toward the base; with an index from 7 to around 12. Fig. 11.

Narrowly oblanceolate – inversely narrowly lanceolate. Fig. 11.

Narrowly obovate – inversely narrowly ovate. Fig. 11.

Narrowly ovate – egg-shaped in outline, broader at the base, two-dimensional;

with an index about 1.6–3. Fig. 11.

Nectary – a nectar-producing structure or gland.

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26

Fig. 9. Orchid morphology (subfamily Vandoideae).

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.


Turczaninowia 2008, 11(1) : 5–168

Fig. 10. Main types of pollinaria and pollinium stalk in orchids: a, b – sagittal section of column

(a) and pollinarium (b) of flower of Tropidia curculigoides, that illustrate hamulus formation;

c, d – sagittal section of column (c) and pollinarium (d) of flower of Doritis pulcherrima, that

illustrate stipe (tegula) formation; e, f – sagittal section of column (e) and pollinarium (f) of

flower of Dactylorhiza incarnata, that illustrate caudicle formation.

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Node – the point on a stem at which a leaf or bract is attached. Fig. 3.

Non-resupinate flower – the flower with not-resupinate ovary having the lip

turned upwards.

Obcordate – inversely cordate. Fig. 11.

Oblanceolate – inversely lanceolate. Fig. 11.

Obligate – restricted to particular conditions or circumstances, like

achlorophyllous mycotrophic plants. Fig. 1, b.

Oblique – with unequal sides; slanting. Fig. 12.

Oblique bifid – divided into 2 unequal shallow segments, usually at the apex.

Fig. 12.

Oblong – much longer than broad, with nearly parallel sides, rounded at each

end. Fig. 11.

Obovate – inversely ovate. Fig. 11.

Obsolete – wanting or rudimentary.

Obtuse – blunt or rounded at the apex. Fig. 12.

Ombrophilous – adapted to shady habitats of humid rainy forest.

Operculum – see anther cup. Fig. 8.

Orbicular– of a flat body with a circular outline. Fig. 11.

Oval – oval-shaped, not much longer than broad, with nearly parallel sides,

rounded at each end. Fig. 11.

Ovary – the part of the flower that contains ovules and develops into the fruit.

Fig. 3–5, 8.

Ovate – egg-shaped in outline, broader at the base, two-dimensional; with an

index around 1.4–1.6. Fig. 11.

Ovoid – egg-shaped, three-dimensional.

Palea – flat or terete, moveable appendages attached by a filiform base, found

on tepals in some species of Bulbophyllum. Fig. 8.

Pandurate – violin- or fiddle-shaped. Fig. 11.

Panicle – a much branched inflorescence.

Paniculate – branched, branching (of an inflorescence).

Papilla (pl. papillae) – a soft, short, superficial gland or protuberance. Fig. 6.

Papillose – covered with papillae.

Parietal placentation – placentation when ovules are attached to the walls

of the ovary. Fig. 2, b.

Pectinate – comb-like; with close, regularly spaced teeth or hairs, often in a

single row, like the teeth of a comb.

Pedicel – the stalk of a single flower in an inflorescence. Fig. 3, 8.

Peduncle – the stalk bearing an inflorescence or solitary flower. Fig. 3, 5, 6.

Peloric – a more or less radially symmetrical mutant of a species which

normally has zygomorphic flowers.

Peltate – shield-shaped.

Pendent – hanging.

Pendulous – hanging.


Turczaninowia 2008, 11(1) : 5–168

Perianth (tepals) – a collective term for sepals and petals, together.

Petals – in orchids two lateral segments of the inner whorl of perianth (corolla).

Fig. 4, 5, 7, 9.

Petiolate leaf – leaf with distinct petiole and leaf blade.

Petiole – a leaf stalk. Fig. 6.

Pilose – soft hairy.

Pistil – the female reproductive organ of a flower, typically consisting of a

stigma, style and ovary.

Placenta – that portion of the ovary that bears the ovules.

Placentation – the arrangement of ovules and configuration of the placentas

in ovary. Fig. 2, a, b.

Plicate – pleated or folded leaves having several to many major longitudinal

veins and folded in each one. Fig. 2, l.

Pollen masses – see Pollinium.

Pollinarium (pl. pollinaria) – the complete set of pollinia with associated parts,

like viscidium (viscidia) and stipe. Fig. 6–10.

Pollinium (pl. pollinia) – a more or less compact mass of pollen. Fig. 6–10.

Porate pollinium – pollinium with a pore or pores.

Porrect – directed outward and forward.

Prostrate – lying flat on the ground.

Proteranthous – an apical inflorescence produced before the pseudobulbs

and leaves on the same shoot. Fig. 2, g.

Proximal – the part nearest the axis, toward the base.

Pseudobulb – a swollen aerial stem. Fig. 3, 8.

Pseudocopulation – a special type of pollination, in which flowers mimic

female insects and are pollinated by the males when they attempt to copulate with the

flowers.

Pseudopollen – mealy, farinose, pollen-like deposit (originated from highly

modified papillae) on the lip of some species of Eria and Dendrobium.

Pubescent – softly hairy or downy.

Pulvinate – cushion-like.

Pustulate – see pustulose.

Pustulose (pustulate) – having small blisters or pustules.

Pyriform – pear-shaped. Fig. 11.

Quaquaversal – directed or bending in every direction.

Raceme – an unbranched, elongate, indeterminate inflorescence with

pedicellate flowers maturing from the base upwards.

Racemose – having flowers in racemes.

Rachis – the axis of an inflorescence. Fig. 3, 8, 9.

Recurved – curved backwards or downwards.

Reflexed – abruptly bent or turned downwards or backwards.

Reniform – kidney-shaped. Fig. 11.

Resupinate flower – the flower with resupinate ovary having the lip turned down.

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 11. Main types of shapes and margins of orchid leaf and tepal.


Turczaninowia 2008, 11(1) : 5–168

Fig. 12. Main types of apices and bases of orchid leaf and tepal.

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Reticulate – netted.

Retrorse – directed backwards or downwards.

Retuse – shallowly notched at a rounded apex. Fig. 12.

Revolute – rolled back from the margin.

Rhizome – horizontal, underground or surface-resting stem bearing scale

leaves and adventitious roots; in sympodial orchids composed of the bases of successive

shoots. Fig. 3, 6.

Rhombic – diamond-shaped. Fig. 11.

Rhomboidal – quadrangular, nearly rhombic, with obtuse lateral angles.

Root ridge – elongate callus on fleshy rhizome which bears tufts of root hairs

in some rootless orchids like species of Cheirostylis.

Root-stem tuberoid – see tuberoid.

Rosette – a densely clustered spiral of leaves, usually borne near the ground.

Rostellum – highly modified, often beak-like sterile median portion of stigma

lying between the functional part of stigma (stigmas) and stamen. Fig. 6–10.

Rostellar arms – lateral narrow wing-like rostellar extensions in some

representatives of subtribe Goodyerinae. Fig. 6.

Rostrate – beaked.

Rosulate – with the leaves arranged in basal rosettes on very short stem.

Rotund – circular in outline.

Round – see rounded.

Rounded (round, rotund) – with a rounded apex or base. Fig. 12.

Rugose – wrinkled.

Rugulose – somewhat wrinkled.

Saccate – with a conspicuous swelling hollow.

Sagittal section – see longitudinal section.

Sagittate – arrowhead-shaped. Fig. 12.

Saprophyte – plants which obtain nutrients by absorption of organic substances

from the soil (as a rule, indirectly, by means of mycorrhizal symbiotic or mycotrophic

relationships with soil fungi). Fig. 1, b.

Saprophytic – see mycotrophic.

Scale – any thin, flat, scarious structure commonly of leaf origin. Fig. 3.

Scandent – climbing.

Scape – a leafless and flowerless lower part of an inflorescence. Fig. 3, 5–7, 9.

Scarious – thin, dry and membranous.

Scattered – arranged without apparent order.

Sectile – the conditions in which soft, granular pollinia are subdivided into

small packets, which are usually connected by elastic threads.

Secund – arranged on one side of the axis only.

Semicircular – half-round. Fig. 11.

Sepals – in orchids three segments of the outer whorl of perianth (calyx). Fig. 4.

Septum – a partition, like in the spur or sac in many species of subtribe

Aeridinae.


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Serrate – saw-like; toothed along the margin, with the sharp, regular teeth

pointing forward. Fig. 11.

Serrulate – saw-like; toothed along the margin with minute, sharp, regular

forward-pointing teeth. Fig. 11.

Sessile – attached directly, without stalk or petiole.

Sessile leaf – leaf lacking the petiole.

Seta (pl. setae) – a bristle.

Setaceous – bristle-like; with bristles.

Sheath – a leaf-like structure which enfolds a stem, pseudobulb, or young

inflorescence.

Sigmoid – S-shaped.

Spatulate – oblong and attenuated at the base, like a spatula. Fig. 11.

Spicate – arranged in a spike.

Spike – an unbranched inflorescence with sessile or subsessile flowers

maturing from the bottom upwards.

Spur – a slender, tubular or sac-like often nectar-containing projection

commonly formed by the base of the lip. Fig. 6, 7, 9.

Stamen – the male reproductive organ of a flower, consisting of an anther

and filament. Fig. 5.

Staminode (staminodium) – a sterile stamen, often modified in shape and

size. Fig. 4, 5, 7.

Staminodium – see staminode.

Stelidium (pl. stelidia) – a wing- or arm-like lateral appendage on the top of

the column. Fig. 8, 9.

Stigma – the portion of the pistil which is receptive to pollen. Fig. 4–6, 8–10.

Stigma lobes – two completely separated lateral lobes of stigma. Fig. 7.

Stigmatophore – an elongated process on either side of the base of the

column bearing lateral stigmatic lobe. Fig. 7.

Stipe (tegula) – a non viscid band or strap of columnar tissue which connects

the pollinia to the viscidium. Fig. 9, 10.

Stipitate – raised on a stalk.

Style – the slender part of the pistil which connects the ovary with the stigma;

forms a part of the column. Fig. 4, 5.

Subacute – slightly acute. Fig. 12.

Subulate – awl-shaped. Fig. 11.

Sulcate – with longitudinal grooves or furrows.

Sympodial growth – growth in which each new shoot is determinate and

terminates in one season with terminal or lateral inflorescence. Fig. 2, d, e.

Synanthous – an inflorescence when pseudobulb, leaf and apical inflorescence

are produced together. Fig. 2, h.

Synsepal (synsepalum) – a compound organ formed by the union of two

lateral sepals, like in species of Paphiopedilum. Fig. 5.

Tegula – see stipe.

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Tepal index – see index.

Tepals – see perianth.

Terete – cylindric and usually tapering. Fig. 11.

Terminal – the end or apex.

Terrestrial – ground dwelling plants, usually with creeping rhizome, ground

pseudobulbs or subterranean tuber (root-stem tuberoids). Fig. 1, a.

Tessellate – chequered.

Theca (pl. thecae) – a pollen sac or cell of the anther. Fig. 7.

Tomentose – densely covered with matted woolly or short hairs.

Triangular – three-angled, with an index more than 0.85. Fig. 11.

Trichome – a hair or hair-like outgrowth of the epidermis. Fig. 9.

Tridentate – three-toothed. Fig. 12.

Truncate – ending abruptly, as though broken off. Fig. 12.

Trunk epiphytes – see bark epiphytes. Fig. 1, f.

Tuber – in orchids see tuberoid.

Tuberoid (root-stem tuberoid) – a thickened, more or less modified tuber-like

root with a bud and some anatomical stem structures at the base. Fig. 7.

Umbo – a small blunt or rounded protuberance, as on the ventral surface of

staminode in some species of Paphiopedilum. Fig. 5.

Umbonate – having a central, projecting point, like the boss of a shield.

Uncinnate – hook-shaped.

Undulate – waved.

Unguiculate – contracted at the base into a claw. Fig. 12.

Urceolate – pitcher-, vase- or urn-shaped.

Velamen – one or more layers of hygroscopic spongy dead cells on the outside

of a root.

Ventral – refers to the front or inward surface of an organ in relation to the

axis; see adaxial.

Ventricose – swollen or inflated on one side.

Verrucose – warty.

Verticillate – whorled.

Villose – with long, weak hairs.

Vine – see liana.

Viscidiophore – an elongated prolongation of the base of theca (anther canal)

and rostellum side lobes bearing at the apex viscidium. Fig. 7.

Viscidium (pl. viscidia) – a viscid, usually disc-like part of the rostellum joined

to the pollinium, enabling it to adhere to an insect-pollinator body. Fig. 6-10.

Whorl – a circle of three or more leaves or leaf-like organs attached at the

same level on a stem.

Zygomorphic – bilaterally symmetrical.


Turczaninowia 2008, 11(1) : 5–168

ACKNOWLEDGEMENTS

Laboratory studies and work with manuscript and illustrations was supported

by Swiss Orchid Conservation fund of Zurich Foundation for Orchid Conservation

of Swiss Orchid Society. Project – “Orchids of Vietnam, work on monograph

manuscript”, 2005–2007.

We cordially thank authorities of the Institute of Ecology and Biological Resources

of the Academy of Science and Technology of Vietnam and Komarov Botanical

Institute of the Russian Academy of Sciences for comprehensive help in organizations

of all our investigations. Field studies in Vietnam, the results of which are presented in

this paper, were funded by grants from the next organizations:

U.S.A. National Geographic Society – “Flora of Highlands of the South

Vietnam” 1993–2001 years (grants # 5094-93, 5803-96, 6383-98); “Limestone Flora

of Cao Bang Province of northern Vietnam” 1998–1999 years (grant # 6300-98);

“Botanical Inventory of Unexplored Areas in Viet Nam: The North” 1999–2001

(grant # 6733-00); “Exploration of rocky limestone flora and vegetation in Bac

Kan province, northern Vietnam” 2003–2005 (# 7577-04); “Exploration of highland

flora and vegetation in Lai Chau & Son La Provinces, north-western Vietnam”

2006–2007 (#8074-06)

U.S.A. National Science Foundation – “Collaborative Research: A Multi-

Taxa Inventory of Threatened Conservation areas in Viet Nam” 1999–2001 (grant

# DEB-9870231).

Henry Luce Foundation. Vietnam Botanical Conservation Program in

Vietnam. 1999–2006. “Preliminary updated checklist of orchids (Orchidaceae)

of Ba Be National Park”, 2002; “Preliminary updated checklist of orchids (Orchidaceae)

of Bach Ma National Park”, 2003; “Preliminary updated checklist of

orchids (Orchidaceae) of Nui Chua National Park”, 2004; “Preliminary updated

checklist of orchids (Orchidaceae) of Bu Gia Map National Park”, 2005; “Preliminary

Survey of the Flora and Vegetation of Bi Doup – Nui Ba National Park”,

2005; “Preliminary Survey of the Flora and Vegetation of Da Krong Nature Reserve

and allied areas”, 2006.

American Orchid Society. “Exploration of endangered Vietnamese Paphiopedilums”,

years 1996–1997; “Exploration of endangered calcium dependent

orchid flora in inaccessible rocky limestone areas of the North Vietnam”,

years 1999–2000; “Population studies of endemic Paphiopedilum species in northern

Vietnam”, 2001–2002 years; “Discovery of endemic orchid flora in remote

limestone areas of Northern Vietnam”, years 2003–2005; “Exploration of Vietnamese

orchid flora in regions allied to Laos territory”, years 2008–2009.

International Cooperative Biodiversity Groups (ICBG) Grant No.

1-UO1-TW01015-01, through funds from the National Institutes of Health, National

Science Foundation, and the Foreign Agriculture Research Service of the United States

Department of Agriculture, U.S.A., 2000–2005.

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

San Diego County Orchid Society Conservation Committee. “Exploration

of endangered calcium dependent Paphiopedilum species in Northern

Vietnam”, 1999–2000 years.

Fauna & Flora International (Vietnam Program). “The distribution of

Paphiopedilum vietnamense and its current status in the wild”, 2000; “Community-based

Conservation of the Hoang Lien Mountain Ecosystem, Vietnam, Flora

and vegetation survey of Van Ban district, Lao Cai province of northern Vietnam”,

2002; “Preliminary survey of orchids and gymnosperms in Trung Khanh

district, Cao Bang province northern Vietnam”, 2004;

Fauna & Flora International Vietnam Conservation Support Program

and the Ministry of Agriculture and Rural Development, Vietnam. “Pu Luong

– Cuc Phuong Limestone Landscape Conservation Project, Preliminary botanical

survey of primary vegetation in Pu Luong nature reserve”, 2003.

Fauna & Flora International (Vietnam Program) & Counterpart International.

“Preliminary survey of Orchids (Orchidaceae) in Phong Nha – Ke

Bang National Park”, 2005.

WWF Indochina Programme. Green Corridor Project in Thua Thien – Hue

Province, VN085301. “Lowland flora and vegetation. Preliminary survey”, 2005.

Basic Research program in Life Sciences of Viet Nam, # 611001.

“Threatened conifers and cycads of Vietnam”, 2000–2002.

Author cordially thanks Ms. A. Averyanova and Ms. T. Maisak, who were

very helpful in preparation of line drawings, as well as thanks Prof. Phan Ke Loc for

photographs used in Fig. 24, c; 31, f; 36, d; 43, g; 44, d and Dr. H. Koopowiz for

photograph used in Fig. 26, i.


Turczaninowia 2008, 11(1) : 5–168

KEY FOR IDENTIFICATION OF ORCHID GENERA

IN THE FLORA OF VIETNAM

Orchidaceae Juss.

1789, Gen. Pl.: 64 (sub “Orchideae”). – Fam. Apostasiaceae Lindl., 1833,

Nixus Pl.: 22 (sub “Apostasieae”). – Fam. Cypripediaceae Lindl., 1833, l.c.: 22 (sub

“Cypripedieae”). – Fam. Neottiaceae Horan., 1834, Prim. Lin. Syst. Nat.: 50. – Fam.

Vanillaceae Lindl., 1835, Key Bot.: 73. – Fam. Limodoraceae Horan., 1847, Сharact.

Ess. Fam. Reg. Veg.: 44.

Typе: Orchis L.

750–800(158) genera and 20000–25000(1005) species. Cosmopolites, except

extra arid, salted, and polar areas.

1. Plants green, photosynthetic, with normal green leaves, green stem or green roots ....... 2

– Mycotrophic (saprophytic), achlorophyllous leafless plants, without any green parts ... 151

2. Flowers with 2–3 fertile stamens, pollen waxy, amorphous, sticky or powdery, not

forming distinct pollinia ..................................................................................................... 3

– Flowers with 1 fertile stamen, pollen mass commonly forming coherent pollinia ........ 5

3. Leaves thin, plicate; all tepals sub-similar, flowers sub-actinomorphic, with 2–3 stamens;

staminode if present small, inconspicuous; ovary with 3 chambers ............................. 4

– Leaves coriaceous, conduplicate; tepals very unequal, lateral sepals joined, lip deeply

saccate; flowers distinctly zygomorphic, with 2 lateral stamens and large shield-like

median staminode; ovary with 1 chamber .................................................... Paphiopedilum

4. Tepals more than 1.5 cm long, ovate; flowers with 3 stamens; inflorescence simple,

erect ........................................................................................................................... Neuwiedia

– Tepals less than 1.5 cm long, narrowly-ovate to lanceolate; flowers with 2 stamens;

inflorescence normally branched, curved and spreading .................................... Apostasia

5. Mainly terrestrial and lithophytic plants without pseudobulbs; leaves plicate, thin

(not succulent), not articulated at the vase; inflorescence terminal; anther wall not

caducous; pollinia soft, granular or sectile ...................................................................... 6

– Terrestrial, lithophytic and epiphytic plants; shoots sympodial pseudobulbous, or

monopodial thin and rigid; leaves commonly conduplicate, coriaceous or succulent,

usually articulate at the base; inflorescence lateral, rare terminal; anther wall form

caducous cap (operculum); pollinia hard, very rare sectile ............................................ 7

6. Plants with creeping rhizome or underground tubers; anther erect or slightly

incumbent; pollinia apically attached to viscidium; rostellum elongate ....................... 8

– Plants with underground root-stem tuberoids; anther erect, at the base firmly united

with column; pollinia sectile with caudiculae basally attached to viscidium or viscidia,

rare without caudicles; rostellum in form of fold protruding between thecae ............ 28

7. Sympodial plants; pollinia soft, waxy, rare cartilaginous, without stalks or caudiculae,

very rare with stipes or hamulus; anther erect in early stages of ontogeny ................ 38

– Sympodial or monopodial plants; pollinia solid, cartilaginous or bony, commonly

with stipes; anther incumbent in early stages of ontogeny, often strongly deflexed at

maturity ................................................................................................................................. 85

8 (6). Stem erect, with a close, more or less dense fascicle of roots at the base .................... 9

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

– Stem decumbent, rhizome-like, rooting at the nodes, at the apex ascending and bearing

a terminal long stalked inflorescence .............................................................................. 14

9. All normal leaves radical, the stem only with rudimentary leaves and sheaths .......... 10

– Normal leaves distant along all length of the stem, or stem at the middle with 2

opposite leaves ................................................................................................................... 11

10. Flowers less than 5 mm across, resupinate, in dense twisted spike; leaves 3–10,

narrowly-lanceolate to linear ................................................................................. Spiranthes

– Flowers 1.5–2.5 cm across, not resupinate, in loose not twisted spike; leaves 1–2 (3),

ovate ....................................................................................................................... Cryptostylis

11. Stems weak, slender, 1–1.5 mm thick, less than 15 cm tall; at the middle with 2 broadly

cordate opposite leaves 1–2(3) cm wide ...................................................................... Listera

– Stems robust and rigid, more than 1.5 mm thick, more than 15 cm tall; with numerous

distant leaves, rarely with 2 sub-opposite leaves 5–12 cm wide .................................. 12

12. Stems 1.5–3 m tall; inflorescence normally branching; sepals and petals 2–3 cm long

................................................................................................................................ Corymborkis

– Stems less than 1.5 m tall; inflorescence simple; sepals and petals 5–15 mm long ..... 13

13. Flowers, in many flowered spike or in dense heads; sepals and petals white, 0.6–1 cm

long; lip simple narrowly-ovate, boat-shaped ........................................................ Tropidia

– Flowers in lax few flowered raceme; sepals and petals yellow-green or pink-purple;

1.2–1.8 cm long; lip distinctly 3-lobed, cup-shaped .............................................. Epipactis

14 (8). Flowers not resupinate (or partially resupinate), lip commonly turned upwards ....... 15

– Flowers resupinate, lip commonly tuned down ............................................................. 16

15. Leaves uniformly green; lip and column not twisted .............................................. Hetaeria

– Leaves with contrast golden or reddish reticulated veins; lip and column distinctly

twisted .......................................................................................................................... Macodes

16. Spur at the base of lip containing neither glands nor hairs ........................................... 17

– Spur hairy within or with papillae or glands on either internal lateral side .................. 18

17. Spur short and fat, epichile of lip broad triangular acute without any ornaments;

sepals olive-green, petals and lip white ................................................................ Erythrodes

– Spur long and slender, epichile of the lip truncate to slightly bilobed; disc of lip with

two flat calli at the middle and a thick lamella extending to the tip; tepals including lip

white with pink blotches ............................................................................................ Herpysma

18. Spur densely haired within ....................................................................................... Goodyera

– Spur with a single gland or group of papillae on either internal lateral side ................ 19

19. Spur with group of dense numerous, long, fat papillae on either internal lateral side;

lip strongly S-bended; mesochile with two tall conspicuous semicircular longitudinal

keels ........................................................................................................................ Orchipedum

– Spur with a single gland or few papillae on either internal lateral side; lip not strongly

bended; mesochile without keels, rare with 1–2 rather low longitudinal keels ........... 20

20. Apex of the lip not widening into a blade, broadly acute, fleshy .................... Vrydagzynea

– Apex of the lip widened to a flat thin commonly bilobed blade often dentate or fringed

along margin ........................................................................................................................ 21

21. Mesochile of the lip form distinct conspicuous claw as long as epichile or longer ... 22

– Mesochile of the lip very short, appear as constriction between cup-like base of lip

and epichile .......................................................................................................................... 26

22. Claw of lip fringed or toothed along either lateral side .................................................. 23

– Claw of lip straight along lateral margins ........................................................................ 24


Turczaninowia 2008, 11(1) : 5–168

23. Lip spurred, the spur protruding between the lateral sepals; stigma lobes 2, on either

side of base of rostellum; leaves usually deep velvety green, brown or black with

silvery, golden or reddish net of veins ............................................................ Anoectochilus

– Lip with small, globular, saccate base, more or less enclosed by the bases of the

lateral sepals; stigma entire; leaves commonly uniform green, rare plant aphyllous .....

............................................................................................................................... Odontochilus

24. Tiny plants 4–6 cm tall with leaves about 1 cm long; inflorescence 1–2-flowered;

column short, not twisted; stigmas on short process ......................................... Myrmechis

– Plants normally larger than 6 cm tall with leaves much longer than 1 cm; inflorescence

normally few to many-flowered; column twisted or not twisted; stigmas sessile ...... 25

25. Column and lip distinctly twisted; mesochile without longitudinal keels .............. Ludisia

– Lip and column not twisted, rarely slightly twisted; mesochile with 1–2 more or less

conspicuous longitudinal keels ............................................................................. Rhomboda

26 (21). Leaves black; epichile 3-lobed, brightly yellow, with 4 close callosities on the disc ....

..................................................................................................................................... Zeuxinella

– Leaves green to gray-green; epichile 2-lobed, flat, without any ornament ................. 27

27. Sepals free; epichile not toothed or fringed along margin; plant with creeping stem

rooting at nodes ............................................................................................................ Zeuxine

– Sepals connate on half way of their length, epichile distinctly toothed or fringed

along margin; plants with succulent creeping rootless articulate rhizome ... Cheirostylis

28 (6). Tiny plants less than 6 cm tall with 1 cordate leaf and single large flower; lip with 2

short spurs; lateral sepals filiform, much narrower than large broad median sepal;

pollinia without caudicles ........................................................................................... Corybas

– Plants regularly taller than 6 cm, with 1 to many leaves; inflorescence with few to

many flowers, rare with 1 flower; lip with 1 spur or sac, rarely spurless; all sepals

ovate to elliptic, subsimilar; pollinia with caudicles and basal viscidia ....................... 29

29. One or two stigmas distinctly concave ........................................................................... 30

– Stigmas convex or stalked, rarely hardly concave ........................................................ 32

30. Many-leaved stem with distant leaves; floral bracts larger than flowers, similar to

normal leaves; tuberoids palmate or attenuate ............................................ Brachycorythis

– Plant with 1–2(3) radical leaves; floral bracts inconspicuous, shorter than leaves;

tuberoids attenuate to globose ........................................................................................ 31

31. Lip deeply 3-lobed; plant with 1–2(3) ovate to narrowly ovate, uniformly green leaves;

stigma divided into 2 lobes; rostellum very small inconspicuous; tuberoids attenuate

.................................................................................................................................. Amitostigma

– Lip entire or very indistinctly trilobe; plant with 1(2) circular to broadly ovate, purpleviolet

variegated leaves; stigma entire, V-shaped; rostellum large elongate, hornshaped,

curved upwards; tuberoids globose ....................................................... Hemipilia

32 (29). Rostellum very large, horn-shaped, turned upward, as high as or higher than anther ......... 33

– Rostellum much smaller, inconspicuous, shorter than half of column length, sometimes

hardly visible ....................................................................................................................... 34

33. Plants 6–10 cm tall with 1–2 narrowly-lanceolate leaves; inflorescence normally with

1(2) flowers; flowers white; lip entire or indistinctly 3-lobed ............................. Diplomeris

– Plants normally taller than 10 cm with 3–10 broadly-lanceolate leaves; inflorescence

normally with (1)3–20 flowers; flowers purple, orange-red to deep yellow; lip distinctly

4-lobed ....................................................................................... Habenaria (H. rhodocheila)

34. Two stigma-lobes placed on elongated more or less long stigmaphores extending

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

forward from the column base ............................................................................... Habenaria

– Stigma-lobes not freely extending in front of column ................................................... 35

35. Thecas at their base extending from column base; rostellum forming a broad band

above the coherent, hardly prominent stigma-lobes ............................................. Pecteilis

– Neither stigma-lobes nor thecas extending in front of column base ............................ 36

36. Lip ligulate, undivided, at the base with long narrow spur not dilated toward apex;

stigma sessile, with a single hardly concave surface ....................................... Platanthera

– Lip usually 3-lobed; spur if present short saccate or globular; stigma-lobes separate

or hardly coherent .............................................................................................................. 37

37. Lip without spur; 2 large stelidia rise above anther on lateral sides of the column .........

.................................................................................................................................... Herminium

– Lip with short, but distinct spur at the base; stelidia small insignificant, sometimes

hardly visible .............................................................................................................. Peristylus

38 (7). Creeping epiphytic or lithophytic vines with fleshy succulent green stem to 15 m

long; pollinia very soft and mealy ................................................................................ Vanilla

– Stem more or less short, not liana-like; plants sometimes with creeping plagiotropic

woody rhizome up to 2–3 m long; pollinia mealy to hard .............................................. 39

39. Leaves not articulate, circular, broadly cordate or fan-shaped usually appear after

flowering; plants with globular underground tuber; pollinia soft, sectile ............. Nervilia

– Leaves usually articulate, elliptic, ovate and lanceolate to linear; flowering plants

commonly have leaves, rarely plants develop flowers before the leaves appear; plants

rarely have underground tubers; pollinia mealy to hard, sometimes with caudicles ........ 40

40. Pollinia rather hard, 4–8, with or without caudicles; plants commonly epiphytic with

or without pseudobulbs .................................................................................................... 41

– Pollinia rather soft, 8, rarely 4, with distinct, but sometimes quite small caudicles;

plants commonly terrestrial, often with short vertical stem, plicate leaves and lateral

inflorescence ........................................................................................................................ 43

41. Column not distinctly hooded or winged at the apex; plants commonly with

pseudobulbs of several internodes; inflorescence terminal or lateral; pollinia 4-8 with

or without caudicles ........................................................................................................... 42

– Column hooded or winged at the apex around anther; plants with pseudobulbs of 1

internode; inflorescence terminal (sometimes superficially appearing as lateral); pollinia

4, hard, with small caudicles ............................................................................................. 59

42. Flowers with a short or no columnfoot; leaves conduplicate or plicate ...................... 64

– Flowers with a prominent columnfoot; leaves always conduplicate ........................... 67

43 (40). Plants with tall erect stem bearing normal numerous distichous leaves ..................... 44

– Stem creeping, ascending, sometime pseudobulbous, short and flesh, with few subradical

leaves ....................................................................................................................... 48

44. Inflorescence terminal ........................................................................................................ 45

– Inflorescence distinctly lateral .......................................................................................... 46

45. Flowers white, 8–12 cm across, lip entire or hardly 3-lobed, white with yellow-orange

keels; flower bracts large, thin, deciduous, 3–4 cm long; leaves deciduous, thin, with

blue-green waxy surface; lithophytic or epiphytic plant .......................................... Thunia

– Sepals and petals white, with pink-purple tint, flowers 3–8 cm across; lip distinctly 3lobed,

pink to purple, commonly with yellow center; flower bracts small, insignificant

persistent, usually less than 5 mm long; leaves persistent, rigid, not waxy; terrestrial

or lithophytic plant .................................................................................................... Arundina


Turczaninowia 2008, 11(1) : 5–168

46. Stem at the base inflated, fleshy, pseudobulbous; lip elastically hinged to the column

and connected with column base with 2 elastic flanges .................................. Plocoglottis

– Stem at the base not inflated; lip movably or firmly joined to columnfoot or column

base, not connected with column with elastic flanges .................................................. 47

47. Lip spurred; inflorescence normally with 2–4(6) flowers; flowers purple-violet, 5–

8cm across ....................................................................................................................... Phaius

– Lip without spur or sac at the base; inflorescence normally with more than 10 flowers;

flowers white to yellow-greenish, less than 3 cm across ..................... Cephalantheropsis

48 (43). Stem fleshy, creeping, rooting at nodes with ascending apex and few distant leaves ...... 49

– Stem pseudobulbous, short and flesh, with few sub-radical leaves ............................ 51

49. Flowers not resupinate; lip with short spur, turned upward .................... Nephelaphyllum

– Flowers resupinate; lip with long slender spur or spurless, turned down .................. 50

50. Lip with a long slender spur; inflorescence 1-flowered ..................................... Hancockia

– Lip spurless; inflorescence normally many-flowered ................................. Mischobulbоm

51 (48). Plants with underground tuberous rhizome ................................................................... 52

– Stem pseudobulbous, green, short and flesh, sometimes covered with leaf sheaths ....... 55

52. Flowers purple-violet, not resupinate; sepals connate, forming a narrow tube held at

right angle to the ovary ...................................................................................... Anthogonium

– Flowers white, pinkish to purple-violet and yellow-orange, resupinate; sepals free .... 53

53. Inflorescence terminal; leaves elliptic to ovate ............................................................ Bletia

– Inflorescence lateral; leaves linear to lanceolate, rare narrowly elliptic ...................... 54

54. Inflorescence normally many-flowered, dense, secund spike; flowers not widely

opening, less than 1 cm across, light pinkish; each growth normally with 1(2) linear

leaves ...................................................................................................................... Pachystoma

– Inflorescence normally few-flowered, all-faced, loose raceme; flowers widely opening,

normally more than 1 cm across, pink-violet or yellow; each growth normally with

several lanceolate to narrowly elliptic leaves .................................................. Spathoglottis

55 (51). Pseudobulbs consist of 1 internode ............................................................................... 56

– Pseudobulbs consist of 2 or more internodes ................................................................ 57

56. Pseudobulbs 2-leaved; leaves sessile with broad base; inflorescence, flower bracts,

pedicel, ovary and sepals outside brown hairy ......................................................... Eriodes

– Pseudobulbs 1-leaved; leaves petiolate with long distinct petiole, all plant hairless ....

............................................................................................................................................ Tainia

57. Each growth with 1–2 leaves; sepals connate, forming broad urn-like tube; lip mobile

on a large columnfoot .................................................................................. Acanthephippium

– Each growth usually with more than 2 leaves; sepals free; lip not mobile, firmly attached

to the column base ............................................................................................................. 58

58. Column margins over nearly their whole length connate with base of lip .......... Calanthe

– Lip only connate with column at or near its base ...................................................... Phaius

59 (41). Lip at base flat or slightly concave .................................................................................. 60

– Lip at base saccate ............................................................................................................. 62

60. Lip at base S-curved in lateral view; sepals and petals about equal width .......... Panisea

– Lip at the base more or less flat, not S-curved; petals usually narrower than sepals ..... 61

61. Lip entire to obscurely lobed; bulbs and leaves short-living annual; leaves thin,

herbaceous, deciduous; inflorescence usually 1-flowered; lip at base convolute around

base of column ............................................................................................................... Pleione

– Lip usually distinctly 3-lobed; bulbs and leaves long-living perennial; leaves rigid,

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

coriaceous, not deciduous; inflorescence usually few to many-flowered; lip not

convolute around the column ............................................................................... Coelogyne

62 (59). Flowers large, tepals about 4 cm long; sepals with saccate, concave base; lip funnel

shaped, side lobes large, folded around the column .............................................. Neogyna

– Flowers small or of medium size, tepals less than 2 cm long; sepals at the base more o

less flat; lip side lobes small, not folded around the column ........................................ 63

63. Column long, narrow, dilated at the apex; lip distinctly divided into a deeply concave

hypochile with two high lateral lobes and a flat entire epichile, strongly reflexed

down; epichile and hypochile separated with transversal bilobed band; floral bracts

deciduous .................................................................................................................. Otochilus

– Column short, thick, not distinctly dilated toward the apex; lip not distinctly divided

into hypochile and epichile; lip have no transversal bands; floral bracts deciduous or

persistent ................................................................................................................... Pholidota

64 (42). Leaves distichous, laterally compressed, fleshy ........................................................... 65

– Leaves dorsiventral, flat and thin .................................................................................... 66

65. Leafy stems close together ....................................................................................... Oberonia

– Leafy stems spaced on creeping rhizome ..................................................... Hippeophyllum

66. Column long, slender, commonly longer than broad; flowers resupinate, very rare not

resupinate, lip turned down, very rare upwards, narrowing to the base, without auricles

........................................................................................................................................... Liparis

– Column short, thick, commonly as long as broad; flowers always not resupinate, lip

turned upwards, with broad auriculate base ............................................................. Malaxis

67 (42). Pollinia 4 ............................................................................................................................... 68

– Pollinia 6 or 8 ...................................................................................................................... 76

68. Shoots clustering into dense tufts, stems slender, not pseudobulbous, commonly

less than 10 cm tall, with many close distichous leaves usually less than 1 cm long;

inflorescence terminal, less than 1 cm long, few-flowered with small campanulate

flowers 2–3 mm across; pollinia clavate .............................................................. Podochilus

– Shoots commonly not clustering into dense tufts, stems commonly swollen (at least

in some internodes), pseudobulbous, rare slender, reed-like, normally taller than 10cm;

or short pseudobulbous, consist of one swollen fleshy internode, distant on creeping

woody rhizome; leaves rare less than 1 cm long; inflorescence lateral or sub-terminal

commonly longer than 1 cm, 1- to many-flowered; flowers campanulate to well-opening,

commonly larger than 3 mm across; pollinia not clavate ............................................... 69

69. Pollinia without stipes ........................................................................................................ 70

– Pollinia with stipes .............................................................................................................. 74

70. The non rhizomatous part of sympodial shoots consisting of several to many

internodes, usually fleshy, pseudobulbous (rare thin, reed-like), clustering on short

rhizome .................................................................................................................................. 71

– The non rhizomatous part of sympodial shoots fleshy, pseudobulbous (sometimes

small indistinct), consisting of 1–2 internodes, commonly distant on more or less long

creeping rhizome .................................................................................................................. 72

71. Basal part of non rhizomatous sympodial shoot thin, woody, rod-like, often branching

with fleshy apical internodes forming 1-leaved pseudobulbs; leaves not sheathing at

the base; inflorescence 1(2)-flowered; flowers lasting few morning hours ............ Flickingeria

– The non rhizomatous sympodial shoot not distinctly differentiated into thin and

fleshy portions, or have fleshy base of few swollen internodes, rarely branching,


Turczaninowia 2008, 11(1) : 5–168

normally with several to many leaves sheathing at the base; inflorescence usually

with more than 2 flowers; flowers commonly lasting more than 1 day ........... Dendrobium

72. Inflorescence appears terminal, from the apex of pseudobulb; lip not mobile, not

hinged at base; menthum rather spur-like .......................................................... Epigeneium

– Inflorescence distinctly lateral, from the rhizomatous part; lip mobile, movably hinged

to columnfoot; menthum rather saccate ......................................................................... 73

73. Operculum with conspicuous horn at apex; sepals more or less equal size and form,

spreading ............................................................................................................................ Trias

– Operculum without horn at apex; lateral sepals commonly different in size and shape

from median sepal, spreading or not ............................................................... Bulbophyllum

74 (69). Stipe very large and massive, as broad as long, with broad apex which occupies

whole clinandrium; inflorescence with 1 hardly opening flower ....................... Hamularia

– Stipe narrow and slender, much longer than broad, placed in furrow in front of column

apex; inflorescence many-flowered, rare with 1–2 widely opening flowers ................ 75

75. Petals fascinated to the basal portion of distally naked columnfoot; columnfoot longer

than column; lip with large falcate erect auricles at base; pollinia with 1 common stipe

................................................................................................................................... Monomeria

– Petals fascinated along the whole edge of the columnfoot; columnfoot of equal length

or shorter than column; lip without conspicuous auricles at base; pollinia with 2

stipes .............................................................................................................................. Sunipia

76 (67). Pollinia 6; lip joined to columnfoot, with tall conspicuous appendage on upper surface

toward the base; stem not pseudobulbous with numerous distichous leaves ..... Appendicula

– Pollinia 8; lip not firmly joined to columnfoot, without tall appendage on upper surface;

stem pseudobulbous or not ............................................................................................. 77

77. Lip with a sac-shaped hypochile separated from epichile by transversal ridge; stem

not pseudobulbous, rod-like with numerous internodes and distichous grass-like

leaves; flowers very small, less than 2 mm across, in dense terminal head ...... Agrostophyllum

– Hypochile not separated from epichile by a transversal ridge; stem pseudobulbous or

not; flowers commonly larger than 2 mm across, not in dense terminal heads ........... 78

78. Pollinia joined by a common long thin caudicle ............................................................. 79

– Pollinia not joined by a common long thin caudicle ...................................................... 80

79. Plant with flat normally 2-leaved pseudobulbs, rare stem not pseudobulbous with few

sub-radical distichous leaves; flowers hardly opening, sub-cleistogamous; lip at the

base with a longitudinal thickening; operculum vertical behind column, beaked; column

without columnfoot ..................................................................................................... Thelasis

– Plant with several sub-radical distichous leaves; flowers campanulate; lip without

thickening; operculum horizontal on top of column, not beaked; column with small,

but distinct columnfoot ............................................................................................. Phreatia

80. Column with 2 vertical arms or lobules bearing stigmas; stem thin not pseudobulbous

.................................................................................................................................. Ceratostylis

– Column without vertical arms or lobules bearing stigmas; stem pseudobulbous or not

.................................................................................................................................................. 81

81. Sepals free; flowers widely opening or campanulate .................................................... 82

– Sepals connate, forming tube; flowers hardly opening ................................................ 84

82. Young leaves densely haired ................................................................................ Trichotosia

– Leaves not hairy ................................................................................................................. 83

83. Lip movably joined to a columnfoot; column long, forward curved; columnfoot

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

distinctly broader than column, in front with large fleshy cushion .................. Callostylis

– Lip not very movable, joined to a columnfoot; column short, not strongly curved;

columnfoot not much broader than column, without conspicuous cushion .......... Eria

84 (81). Pseudobulbs elongate, 4–10 cm tall, normally with 3–5 leaves; inflorescence 3–15 cm

long, of many distichous flowers ....................................................................... Cryptochilus

– Pseudobulbs flat, less than 1 cm tall, normally with 2 leaves; inflorescence less than

0.5 cm long, normally with 1–2 flowers ........................................................................ Porpax

85 (7). Sympodial plants ................................................................................................................. 86

– Monopodial plants .......................................................................................................... 98

86. Lateral sepals free or partially merged into a synsepalum ......................................... 87

– Lateral sepals merged completely into a synsepalum ................................... Acriopsis

87. Stem pseudobulbous; pseudobulbs 1-leaved, consist of 1 internode; column much

curved, apically with 2 forwards curved horns .............................................................. 88

– Stems usually of several internodes, pseudobulbous or not, with 1 to several leaves;

column straight or slightly curved, apically without large, conspicuous horns ........ 89

88. Lip 2-lobed at apex with small insignificant median dent; column S-curved; on the

upper side of columnfoot a narrow nectary opening at base of lip; pollinia 2; operculum

semi globular ............................................................................................................. Thecostele

– Lip at the apex acute, entire; column bent forwards, but not S-curved; columnfoot

hollow with an entrance near the articulate base of lip; pollinia 4; operculum conical

....................................................................................................................................... Thecopus

89. Plants develop neither pseudobulbs nor tubers, stem thin, not pseudobulbous ...... 90

– Plants develop tubers, corms or pseudobulbs of several internodes, or pseudobulbous

stems often covered by close leaf sheaths ..................................................................... 92

90. Terrestrial grass-like plants with erect or climbing stem up to 1 m long; lip densely

haired along midvein ............................................................................................... Dipodium

– Epiphytes with stem less than 20 cm tall, not grass-like; lip not haired ....................... 91

91. Leaves linear, grass-like, laterally compressed or strongly conduplicate; inflorescence

dense, not branching distichous spike; flowers resupinate, lasting few hours ... Bromheadia

– Leaves lanceolate to ovate, hardly conduplicate or flat; inflorescence lax, often

branching all-facing raceme or panicle; flowers not resupinate, long lasting ......... Polystachya

92 (89). Column at front edge with small horizontal triangular or obtuse wings; anther with 2

pollinia ................................................................................................................................... 93

– No wings at front of column; anther with 2 cleft or 4 pollinia ........................................ 95

93. Columnfoot simple, without nectary; lip without claw ...................................... Diglyphosa

– Columnfoot with a saccate nectary or a spur-like nectary formed with connate bases

of lateral sepals; lip at base narrowing into a claw ......................................................... 94

94. Columnfoot short, fat, with a saccate nectary opening at base of column between

vertical oblong cushions, lateral sepals not connate at base .................... Chrysoglossum

– Columnfoot long forming a slender spur-like menthum together with connate bases

of lateral sepals .......................................................................................................... Collabium

95 (92). Stem inflated of many internodes, more or less pseudobulbous, usually covered by

dense leaf sheaths; anther with 2 cleft pollinia, or pollinia 4 in pairs; pollinarium

without stipes, viscidium large, broad with truncate front edge ..................... Cymbidium

– Plants with tubers, corms or epigeous corm-like pseudobulbs of a few internodes;

anther with 2 cleft pollinia or pollinia 4 in pairs; pollinarium with short strap-shaped

stipes, viscidium rather large, sometimes forming a single structure with the stipe ......... 96


Turczaninowia 2008, 11(1) : 5–168

96. Raceme nodding down ............................................................................................ Geodorum

– Raceme normally straight and erect ................................................................................ 97

97. Flowers newer open widely; tepals narrowly lanceolate, free; column narrow, long

and straight without columnfoot; anther with 4 pollinia ..................................... Cremastra

– Flowers normally widely opening; sepals and petals broadly-lanceolate to broadlyovate;

lateral sepals often connate at the base forming small or large spur-like menthum;

column with short, but distinct columnfoot; anther with 2 pollinia cleft at the base ......

....................................................................................................................................... Eulophia

98 (85). Anther with 4 more or less equal globular pollinia free from each other ...................... 99

– Anther with 2, sometimes more or less divided pollinia .............................................. 103

99. Very small, normally leafless epiphytes with flat ribbon-like green roots; stem usually

1–3 mm tall; flowers lasting few hours ........................................................... Taeniophyllum

– Plants with green leaves; stem much taller than 3 mm; flowers lasting few days ..... 100

100. Small canopy epiphytes; numerous leaves less than 2 cm long; inflorescence very

short, 2–4 mm long; flowers less than 5 mm across, subsessile ................................. 101

– Large terrestrial, lithophytic or epiphytic plants; leaves few, much longer than 2 cm;

inflorescence 10–30 cm long; flowers more than 5 mm across .................................... 102

101. Leaves laterally compressed, equitant, closely placed along stem; stems 4–8(12) cm

tall; inflorescence of 2 white flowers .................................................................. Microsaccus

– Leaves dorsiventral, sometimes fleshy, succulent, more or less distant; stems 10–

30cm long; inflorescence of 1–4 green flowers .................................................. Adenoncos

102. Terrestrial or lithophytic rosulate plants; stem 2–5 cm long; all leaves radical, close

each other; inflorescence erect, stout; flowers 1.5–2.5 cm across ........................... Doritis

– Epiphytic plants with long pendent stem to 1 m long; leaves distant on 3–6 cm;

inflorescence erect to pendulous, slender; flowers 4–6.5 cm across ................ Esmeralda

103 (98).Anther with 2 pollen-masses each completely divided in more or less unequal semiglobular

free halves ............................................................................................................ 104

– Pollen-masses more integrated ......................................................................................... 128

104. Column without columnfoot ............................................................................................. 105

– Column with columnfoot, sometimes small but always quite distinct ....................... 123

105. Lip firmly adnate to column, not movable ..................................................................... 106

– Lip movable .......................................................................................................................... 122

106. Hypochile sometimes somewhat concave, no spur or sac at base of lip ................... 107

– Lip with distinct spur or sac at the base ........................................................................ 108

107. Stem up to 1.5 m long with leaves 0.5–1 m long and 6–10 cm wide; flowers very fleshy,

4–6 cm across; lip much shorter than median sepal, obtuse .............................. Vandopsis

– Stem up to 12 cm long with leaves 4–10 cm long and 0.8–2 cm wide; flowers not

particularly fleshy, 1 cm across; lip longer than median sepal, forked at the apex into

2 long teeth ............................................................................................................... Diploprora

108. Spur without longitudinal septa inside .......................................................................... 109

– Spur with a distinct longitudinal median septum inside ............................................. 117

109. Spur inside without calli on back-wall ........................................................................... 110

– Spur inside with different ornaments on back-wall ..................................................... 114

110. Stipes linear nearly 4 times as long as diameter of pollinia; epichile fringed to dentate

............................................................................................................................... Ornithochilus

– Stipes around twice as long as diameter of pollinia; epichile more or less entire along

margin .................................................................................................................................... 111

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

111. Lip with transversal ridge or rugose callosities at the base of epichile ...................... 112

– Lip without transversal ridge or callosities on epichile ............................................... 113

112. Large plants with stem up to 1 m tall and leaves 20–30 cm long; hypochile sac-like,

hairy inside ..................................................................................................................... Acampe

– Small plants with stem up to 20 cm tall and leaves 5–10 cm long; hypochile spur-like,

not hairy inside ..................................................................................................... Smitinandia

113. Large plants with climbing stem normally longer than 50 cm; leaves coriaceous, flat,

oblong; flowers red or yellow, 2–5 cm across; lip much shorter than median sepal ........

................................................................................................................................... Renanthera

– Small plant with pendant or creeping stem normally less than 20 cm long; leaves

fleshy, lanceolate to sub-terete; flowers white to purple, less than 5 mm across; lip as

long as or longer than median sepal ................................................................. Schoenorchis

114 (109). Leaves terete; spur back-wall bears inside Y-shaped callus .................... Cleisostomopsis

– Leaves dorsiventral; spur back-wall inside bears a rising tongue ............................. 115

115. Rising tongue on the lip usually hairy, placed close to entrance to spur .................. 116

– Back-wall rising tongue glabrous, placed deep in spur ................................ Pomatocalpa

116. Inflorescence very short, less than 1 cm long, commonly 1–2-flowered; flowers less

than 1.5 cm across ................................................................................................ Trichoglottis

– Inflorescence normally 10–25 cm long; commonly bears more than 2 flowers; flowers

more than 1.5 cm across ...................................................................................... Staurochilus

117 (108). Vine-like plant with climbing or hanging stem and numerous ovate leaves usually

less than 3 cm long; column with 2 long thin glabrous stelidia incurved above apex;

inflorescence very short, less than 1 cm long, 1–2(3)-flowered ................... Pelatantheria

– Stem erect, rare vine-like and ascending, leaves usually longer than 3 cm; column

without large stelidia; inflorescence longer than 1 cm with few to many flowers ..... 118

118. Floral bracts large, longer than flowers; bracts, ovary and flowers densely pubescent

.................................................................................................................................. Cleisomeria

– Floral bracts small insignificant, shorter than flowers; bracts, ovary and flowers

glabrous, rarely papillose .................................................................................................. 119

119. Rostellum projection narrow, very long, twice longer than broad of operculum, supports

a thin linear stipe, to 9 times as long as diameter of pollinia ........................................ 120

– Rostellum projection shorter; stipe usually shorter than 9 diameters of pollinia ..... 121

120. Vine-like plant with climbing or hanging stem commonly 15–30 cm long, rooting along

all its length; leaves distant, coriaceous; rostellum projection distally upwards turned,

stipe linear on all its length ...................................................................................... Micropera

– Plants with short erect stem, usually up to 10 cm tall, rooting at the base, leaves

fleshy, distinctly distichous, arranged into 2 dense rows; rostellum projection distally

curved down, stipe linear broadening to the apex ........................................... Stereochilus

121. Clinandrium top with a rising dorsal longitudinal ridge with a furrow along edge

bearing long linear stipe and dorsally placed pollinia .................................... Sarcoglyphis

– Column without conspicuous dorsal ridge at the top; pollinia not placed dorsally;

stipe very variable in form .................................................................................... Cleisostoma

122 (105). Sepals and petals broadly ovate; stem erect, straight, semi-woody; leaves usually 3–

5 cm broad; lip broadly spatulate at the apex; epichile with high median keel ...... Hygrochilus

– Sepals and petals narrowly-oblong, spatulate toward the apex; stem slender, often

climbing, vine-like; leaves less than 3 cm broad; lip acute at the apex; epichile with

low median keel ............................................................................................................ Arachnis


Turczaninowia 2008, 11(1) : 5–168

123 (104). Lip without distinct spur or sac, but hypochile often more or less concave; leaves

dorsiventral, usually thin, or lacking .............................................................................. 124

– Lip with distinct spur or sac; leaves dorsiventral, usually fleshy or terete ............... 125

124. Stipe short and broad, shorter than diameter of pollinia; plants with normal leaves;

stem usually longer than 1 cm, rooting along all its length ............................ Thrixspermum

– Stipe linear to strap-shaped, about twice as long as diameter of pollinia; leaves usually

lacking; stem usually less than 1 cm long with densely clustering numerous roots ......

................................................................................................................................. Chiloschista

125. Spur of lip with longitudinal septum .................................................................. Cleisostoma

– Spur of lip without longitudinal septum ........................................................................ 126

126. Leaves terete, usually purple ........................................................................... Rhynchogyna

– Leaves dorsiventral, usually green, sometimes with purple tint ................................ 127

127. Stem short, normally less than 2 cm long; leaves thin, ovate, 3–6 cm wide; flowers

more than 1 cm across, in lax few flowered inflorescence normally longer than 5 cm .....

..................................................................................................................................... Kingidium

– Stem 4–20 cm long; leaves lanceolate-oblong, fleshy, less than 1 cm wide; flowers less

than 1 cm across in dense inflorescence 3–5 cm long .................................. Cleisocentron

128 (103). Anther with 2 pollinia, more or less but not completely cleft or split ........................ 129

– Pollen-masses more integrated ......................................................................................... 139

129. Flowers without columnfoot, or columnfoot very indistinct ...................................... 130

– Flowers with distinct, but sometimes short columnfoot ............................................. 135

130. Inflorescence lax raceme, commonly with 2–5 flowers, flowers commonly more than

1.5 cm across ....................................................................................................................... 131

– Inflorescence dense spike with numerous flowers; flowers commonly less than 1.5 cm

across .................................................................................................................................... 132

131. Epichile 2-lobed, truncate or obtuse, more or less flesh or rigid, with entire margin;

stipe short and broad ....................................................................................................... Vanda

– Epichile entire, broad, semicircular, thin, denticulate along margin; stipe narrowlylanceolate

............................................................................................................ Christensonia

132. Flowers 1–1.5 cm across; robust plants with erect semi-woody stem; leaves large,

very rigid, normally 20–40 cm long often with many light-colored veins .... Rhynchostylis

– Flowers much less than 1 cm across; plants smaller with erect or pendent stem; leaves

not particularly rigid, normally less than 15 cm long, uniformly green ...................... 133

133. Tiny canopy epiphytes; leaves normally 2–5 cm long; flowers 2–3 mm across ... Saccolabiopsis

– Medium size epiphytes; leaves normally longer than 5 cm; flowers more than 4 mm

across .................................................................................................................................... 134

134. Long stemmed plants with distant leaves; stipe lanceolate to spathulate, slightly

dilated and curved at apex; spur distinctly inflated at the apex ......................... Robiquetia

– Rather short stemmed plants with few, more or less close leaves; stipe S-shaped,

rising above and behind pollinia, which are separated by a vertical longitudinal stipe

lamella; spur not inflated at the apex .......................................................................... Uncifera

135 (129). Leaves terete; stem vine-like, slender, climbing ............................................ Papilionanthe

– Leaves dorsiventral; stem not vine-like, more or less erect ........................................ 136

136. Mid-lobe of epichile well developed, large, flat, cordate, triangular or oblong;

hypochile flat, concave or form conspicuous forward curved spur .......................... 137

– Mid-lobe of epichile reduced in form of small fleshy dent, spur or sac placed centrally

on the lip .............................................................................................................................. 138

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

137. Hypochile flat or slightly concave; epichile at the base with high keels, teeth or

thread-like bifurcate appendages; inflorescence with few distichous flowers ... Phalaenopsis

– Hypochile form conspicuous forward curved spur; epichile flat without ornaments;

inflorescence with numerous all-faced flowers .......................................................... Aerides

138. Columnfoot as long as or longer than column; rachis of inflorescence not swollen

along whole its length; stem longer than 2 cm, bearing normally several leaves ......... Pteroceras

– Column much shorter than column; rachis of inflorescence distinctly swollen toward

the apex; stem very short, less than 2 cm long, bearing normally 1–3 leaves .... Brachypeza

139 (128). Anther with 2 porate pollinia ........................................................................................ 140

– Anther with 2 whole pollinia, not cleft, split or porate ................................................. 147

140. Lip movable; columnfoot as long as or longer than column; lateral sepals attached

with their bases to the basal portion of distally naked columnfoot; spur entrance

partially closed by a horizontal plate arising from the back-wall of the lip .... Cryptopylos

– Lip not movable; columnfoot shorter, rarely as long as or longer than column; lateral

sepals attached to columnfoot along whole its length; spur entrance free .............. 141

141. Leaves terete, very rigid; inflorescence very short, commonly less than 1 cm long ................ Luisia

– Leaves dorsiventral, sometimes fleshy conduplicate to sub-terete, but not particularly

rigid; inflorescence commonly longer than 1 cm ......................................................... 142

142. Small canopy epiphytes with leaves normally less than 6 cm; mid-lobe of the lip large,

terete, conical at the apex, fleshy, furrowed and concave toward the base; flowers

distichous, opening in succession and lasting few hours ............................... Biermannia

– Medium size epiphytes with leaves commonly longer than 6 cm; mid-lobe of the lip

more or less flat, not terete or fleshy; flowers all-way faced, lasting few days ......... 143

143. No distinct rising side-lobes on the sac-like hypochile, usually just a fleshy rim joining

in front of sac entrance ..................................................................................................... 144

– Distinct, but sometimes small, side-lobes rising from lateral sides of hypochile which

forms a saccate to cylindrical spur ................................................................................. 145

144. Inflorescence umbel-like, less than 3 cm long; epichile broadly fastened in front of the

rim connecting edges of the hypochile, usually crescent-shaped, commonly shorter

than broad; the edges of the stigma not swelling ............................................. Gastrochilus

– Inflorescence dense raceme much longer than 3 cm; epichile narrow, triangular,

concave to the base connecting by lateral sides the edges of the hypochile; the

edges of the stigma concavity much swelling into protruding ring-like structure .............

.......................................................................................................................... Eparmatostigma

145. Epichile thin, much broadening from a narrow base; leaves strongly conduplicate,

semi-terete; inflorescence lax few-flowered raceme, normally longer than 15 cm. ...............

.............................................................................................................................. Holcoglossum

– Epichile more or less fleshy, ligulate-oblong or broadly cuneate, not narrowing toward

the base; leaves distinctly dorsiventral, very rare sub-terete; inflorescence dense,

many-flowered spike, normally shorter than 15 cm ..................................................... 146

146. Epichile ligulate-oblong, more or less flat; lip side-lobes rather straight, no transversal

callus at the base of epichile; stipe broad and rather short, twice longer than diameter

of pollinia .............................................................................................................. Ascocentrum

– Epichile broadly-cuneate, concave; lip side-lobes strongly incurved, closing the

entrance of the spur, at the base of epichile large fleshy transverse callus connecting

the front-edges of side-lobes, curving inwards; stipe narrow, longer than twice diameter

of pollinia .......................................................................................................... Ascocentropsis


Turczaninowia 2008, 11(1) : 5–168

147 (139). Column without distinct columnfoot; stipe linear, much broadening at apex, 3-5 times

as long as diameter of pollinia ........................................................................................ 148

– Column with distinct columnfoot; stipe ovate to broadly-lanceolate, not much

broadening at apex, as long as or few longer as diameter of pollinia .......................... 149

148. Lip side-lobes very large, flat, ciliate or finely fringed along front margin .... Pennilabium

– Lip side-lobes small in form of insignificant dents, entire along margin ............. Malleola

149. Lip movable, 5-lobed; side-lobes and lateral lobes of epichile long, conspicuous,

erect, longer than broad, mid-lobe very small in form of short fleshy dent ........ Grossourdya

– Lip not movable; epichile short, obtuse, entire, very fleshy, lateral lobes insignificant

or broadly oblong to semi-circular as long as broad ................................................... 150

150. Stem usually 4–8 cm long, erect to suberect, with more or less distant leaves;

inflorescence lax, 5–8 cm long; lip with broadly-oblong or semi-circular side-lobes;

mid-lobe fleshy, broadly conical .................................................................. Parapteroceras

– Stem usually less than 4 cm long, pendulous, with close placed leaves; inflorescence

short, head-like, commonly less than 3 cm long with densely placed flowers opening

in succession; lip very fleshy, with broadly obtuse indistinct lobes .......... Tuberolabium

151 (1). Climbing vines commonly 2–10 m long ......................................................................... 152

– Terrestrial erect herbs commonly less than 2 m tall ...................................................... 153

152. Inflorescence rachis and sepals outside densely pubescent; flowers brightly yellow,

2–3 cm across, column stout ....................................................................................... Galeola

– Inflorescence rachis and sepals outside glabrous; flowers light olive green to dull

pink-whitish, less than 2 cm across, column slender ...................................... Erythrorchis

153. Lateral sepals joined each together for a large part of their length ............................ 154

– Lateral sepals free from one another or only joined at the base ................................. 157

154. Sepals and petals joined into 5-lobed tube, split between lateral sepals; flowers hardly

opening; sepals fleshy, light brownish to brownish-pink; stigma at the base of column;

pollinia 2 ..................................................................................................................... Gastrodia

– Sepals and petals free or joined less than half of their length, newer form tube; flowers

widely opening; sepals and petals very thin, white or blue-violet; stigma near top of

column; pollinia 2 or 4 ....................................................................................................... 155

155. Column without stelidia; flowers white ........................................................... Didymoplexis

– Column near apex with conspicuous long decurved hook-like stelidia; flowers white

or blue-violet ........................................................................................................................ 156

156. Median sepal and petals joined to about half of their length, fused with joined lateral

sepals near their base; columnfoot absent; flowers blue-violet .............. Didymoplexiella

– Median sepal and petals free; columnfoot present; flowers white .... Didymoplexiopsis

157 (153). Lip entire, not lobed ....................................................................................................... 158

– Lip distinctly divided into 2–3 lobes .............................................................................. 160

158. Sepals pubescent or mealy outside; sepals and petals yellow to yellow-brown, lip

pale yellow; plants more or less robust, stem reddish-brown to pinkish-brown, rigid;

fruits juicy, cylindrical, berry-like, dull red-purple; tubers numerous, cylindrical, fingerlike

.................................................................................................................................. Cyrtosia

– Sepals glabrous; sepals and petals white, sometimes with black-violet tips, lip white

with purple tint or purple nerves, sometime with black-violet tip; plants slender, stem

white, soft and weak; fruits dry yellowish-white broadly ovate capsule; tuber one,

ovate to cylindrical-ovate .................................................................................................. 159

159. Sepals and petals white, rare with light pink nerves; lip ovate, white or with light pink

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

center or with light purple spots, at the base with short broad spur ................ Epipogium

– Sepals, petals and lip white, with black-violet tips; lip narrowly ovate to broadlylanceolate,

not spurred ...................................................................................... Stereosandra

160 (157). Flowers with a distinct calyculus beneath perianth; lip apex densely long haired;

stem very brittle, dark brown to black, less than 1 mm thick ........................... Lecanorchis

– Calyculus beneath perianth absent; lip at the apex glabrous or papillose; stem not

particularly brittle, white, yellowish, olive to yellowish-brown, more than 1 mm thick

.................................................................................................................................................. 161

161. Inflorescence yellowish-brown to olive-brown, normally with more than 5 flowers; lip

distinctly divided into hypochile, mesochile and epichile ......................................... 162

– Inflorescence white, normally with 1–3 (5) or more flowers; lip not divided into

hypochile, mesochile and epichile ................................................................................... 163

162. Plants more than 25 cm tall; apex of the lip acute to broadly acute ................ Aphyllorchis

– Plants less than 25 cm tall; apex of the lip distinctly bilobed ........................ Odontochilus

163. Flowers brightly yellow, widely opening; sepals and petals more or less thin, lip sidelobes

flat, reflexed, mid-lobe at the apex broadly truncate; column without stelidia ......

...................................................................................................................................... Vietorchis

– Flowers white with purple tint, hardly opening to campanulate; sepals and petals

fleshy, lip side-lobes upward incurved, mid-lobe broadly acute; column at the apex

with large lateral stelidia .................................................................................................. Yoania

SUBFAMILY APOSTASIOIDEAE REICHENB.

1841, Repert. Herb.: 56 (sub “Apostasieae”). – Fam. Apostasiaceae Lindl.,

1833, Nixus Pl.: 22 (sub “Apostasieae”).

Type: Apostasia Blume

2(2) genera and 15(5) species. Tropical and subtropical Asia to N Australia

and New Guinea.

Neuwiedia Blume, 1834, in Hoeven et De Vriese, Tijdschr. Nat. Geschied. 1:

140; id., 1834, Ann. Sci. Nat. (Paris) 2, 2: 93; De Vogel, 1969, Blumea 17, 2: 313–350;

Chen et Lang, 1986, Acta Phytotax. Sin. 24, 5: 346–348.

Type: N. veratrifolia Blume

Sympodial terrestrial herbs with erect semi-woody unbranched stem, often

with few aerial roots at the base. Leaves numerous, spirally arranged, plicate, the

upper becoming bract-like. Inflorescence sub-terminal, erect, many-flowered, dense

raceme. Flowers pedicellate, resupinate, not opening widely, campanulate, hardly

zygomorphic. Tepals free, sub-similar, lanceolate to narrowly obovate, petals are thinner

in texture, the lip a little broader. Column straight, with 3 fertile stamens, filaments

basally jointed to the style, anthers free. Fruit dry capsule or juicy berry. Seeds with

hard alveolate testa.

6–8(1) species. Mainland Southeast Asia to SW Pacific islands.

1. N. zollingeri Reichenb. f., 1857, Bonplandia 5: 58.

Described from Java (“s.l.”). Type (“Zollinger 2808”) – W.


Turczaninowia 2008, 11(1) : 5–168

Distribution. Mainland Southeast Asia, Java, Sumatra, Kalimantan.

Key to varieties

1. Inflorescence glabrous or with very few scattered hairs ....... 1. N. zollingeri var. annamensis

– Inflorescence densely hairy .............................................................................................. 2.

2. Flowers brightly yellow; ripe fruit white ................................. 2. N. zollingeri var. javanica

– Flowers pure white with yellow central spot on lip; ripe fruit orange to red ........................

...................................................................................................... 3. N. zollingeri var. balansae

1a. N. zollingeri var. annamensis (Gagnep.) Aver., comb. nov. – N. annamensis

Gagnep. 1933, Bull. Soc. Bot. France, 80: 350; Seidenf. 1992, Orch. Indochina:

11; Aver. 1994, Identif. Guide Viet. Orch.: 11; Aver. et Averyanova, 2003, Updated

Checklist Orch. Viet.: 46.

Described from S Vietnam (“Annam: Nui-han-heo, pres Nhatrang...”). Type

(“Poilane 4937”) – P.

Plant up to 40 cm tall, glabrous or with few scattered hairs. Leaves numerous,

thin, close together below inflorescence, petiolate; leaf blade narrowly elliptic to elliptic,

acute, 18–50 by 4–6 cm. Inflorescence 6–15 cm long, bearing 30–60 flowers. Floral

bracts triangular, acuminate, 1–2 by 0.3–0.5 cm, as long as flowers. Flowers yellow.

Tepals lanceolate to narrowly obovate, 15–23 by 4–8 mm, obtuse, ending below apex

in short mucro; petals and lip slightly shorter and wider, dorsally with low keel. Stamens

1.2–1.8 cm long, longer than style; anthers 6–8 mm long. Style straight, 1–1.6 cm long

with roundish stigma. Fruit white juicy berry, 5–8 mm across.

Ecology. Shady broad-leaved evergreen lowland and submontane forests on

granite and gneiss. 900 m. Fl. May. Very rare (DD).

Distribution. Vietnam (Khanh Hoa).

Studied specimens. Nui Han Heo, Nhatarang, Poilane, 4937 (P), Poilane, 6180 (P).

Notes. Closely allied to type variety, from which distinctly differs in large

flowers.

1b. N. zollingeri var. javanica (J.J. Smith) de Vogel, 1969, Blumea, 17, 2:

329. – N. javanica J.J. Smith. 1914, Bull. Jard. Bot. Buitenzorg 2, 14: 5.

Described from Java (“… Artana…”). Lectotype (“J.J. Smith s.n. a. 1900”)– BO.

Plants up to 40 cm tall, all parts more or less densely haired. Flowers bright yellow.

Dorsal keel on petals and lip densely hairy. Ripe fruit white juicy berry. Fig. 14, a-d.

Ecology. Shady broad-leaved evergreen submontane forests on granite and

sandstone. 600–950 m. Fl. March. Rare (LR).

Distribution. Vietnam (Dak Lak, Ninh Thuan). Java, Sumatra, Bali.

Studied specimens. Ninh Thuan, Nui Chua mt HLF 4415 (HN, LE); Dak Lak, Chu

Yang Sin mt, HLF 5479 (HN, LE).

Notes. Ripe berries of this variety spicy-fragrant, sweet and edible.

1c. N. zollingeri var. balansae (Gagnep.) Aver., comb. nov. – N. balansae

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Gagnep. 1933, Bull. Soc. Bot. France, 80: 350. – N. griffithii auct. non Reichenb. f.:

Seidenf. 1992, l.c.: 11; Aver. 1994, Identif. Guide Viet. Orch.: 12; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 46. – N. inae auct. non de Vogel: Seidenf. 1992,

l.c.: 13; Aver. 1994, l.c.: 12; Aver. et Averyanova, 2003, l.c.: 46.

Described from N Vietnam (“Tonkin: mt. Bavi, …”). Lectotype (“n° 2000

(Balansa)”) – P.

Plants up to 40 cm tall, all parts more or less densely. Flowers pure white, lip

with yellow spot in center. Dorsal keel on petals and lip densely hairy. Ripe fruit

orange to red juicy berry. Fig. 13, a; 14, e-g.

Ecology. Shady broad-leaved, evergreen lowland and submontane forests of

sandstone, granite and rocky limestone. 450–1250 m. Fl. May – June. Rare (LR).

Distribution. Vietnam (Dak Nong, Khanh Hoa, Kon Tum; Ninh Binh, Phu

Tho, Quang Ninh).

Studied specimens. Cam Pha, Cong Hoa, Quang Ninh, Tran s.n. (LE, UHN); Ninh

Binh, Cuc Phuong, Duc B 356 (HN), CPNP 5116 (HN), N.M. Cuong, H.L. Quyen, NMC 309

(HN), DDS 11060 (HN), Averyanov et al., HAL 1686 (HN, LE), Duc B 356 (HN), N.T. Hiep, NTH

3056 (HN); Dak Nong Prov., Ta Dung mt, HLF 5567 (HN, LE); Mt. Bavi, Ha Son Binh 800 m,

Balansa, 2000; Song Cao near Nhatrang 800 m, Evrard, 508 (AMES) & Poilane, 3408 (P);

Dakto, Gialai-Kontum, Dao, 230 (HN).

Notes. Closely allied to N. zollingeri var. singapureana (Backer) de Vogel,

from which differs in large pure white flowers. Specimens from Cuc Phuong national

park occasionally have flowers with 4 fertile stamens.

Apostasia Blume, 1825, Bijdr. 6, fig. 39; id. 1825, ibid. 8: 423; De Vogel, 1969,

Blumea, 17, 2: 313–350; Chen et Lang, 1986, Acta Phytotax. Sin. 24, 5: 348–351.

Type: A. odorata Blume

Sympodial terrestrial herbs with erect semi-woody commonly unbranched stem,

with few aerial roots at the base. Leaves few to numerous, spirally arranged, plicate,

with acute to filiform tip. Inflorescence sub-terminal, erect to arching and pendent,

usually branched, many-flowered. Flowers, sessile, not resupinate, opening widely,

hardly zygomorphic. Tepals free, sub-similar, broadly lanceolate, more or less recurved,

with dorsal keel extending apically into acute beak, the lip a little larger and wider.

Column straight or curved, usually with 2 fertile stamens, often with 1 staminode,

filaments short, basally jointed to the style; anthers free; stigma 3-lobed or suborbicular.

Fruit dry cylindrical capsule. Seeds with black hard alveolate testa.

7(4) species. Tropical and subtropical Asia to N Australia and New Guinea.

Key to species

1. Plants 5–12 cm tall; leaves petiolate, leaf blade ovate to elliptic, 1.5–2.5 cm long, 0.5–

1 cm wide ........................................................................................................... 3. A. nipponica

– Plants 20–100 cm tall; leaves sessile, linear to lanceolate, 7–35 cm long, 0.3–2.2 cm

wide ........................................................................................................................................ 2

2. Inflorescence at the base with bunch of numerous sterile bracts; flowers white or

yellow; tepals strongly reflexed right back; staminode absent ...................... 4. A. nuda


Turczaninowia 2008, 11(1) : 5–168

– Inflorescence without bunch of sterile bracts at the base; flowers yellow; tepals

reflexed back or straight; staminode present, joined most of the way to style ............ 3

3. Tepals less than 6 mm long; capsule less than 1.8 cm long; column terete, without

wings below staminode; leaves linear to narrowly lanceolate, commonly to 1.5 cm

wide ....................................................................................................................... 1. A. wallichii

– Tepals more than 6 mm long; capsule longer than 1.8 cm; column more or less flattened,

with 2 wings below staminode; leaves narrowly lanceolate to narrowly elliptic,

commonly 1.5–2 cm wide ..................................................................................... 2. A. odorata

1. A. wallichii R. Br. 1830, in Wall. Pl. As. Rar. 1: 75; Seidenf. 1992, Orch.

Indochina: 13; Aver. 1994, Identif. Guide Viet. Orch.: 13; Aver. et Averyanova, 2003,

Updated Checklist Orch. Viet.: 11.

Described from Nepal (“in valle Nagaliae minora, Noakote dicta”). Lectotype

(“Wallich, 4448”) – K (holo), E (iso).

Plant 15–100 cm tall. Leaves numerous, close, linear to narrowly lanceolate,

12–35 by 3–15 mm. Inflorescence, simple to many branched, many flowered. Flowers

pale yellow. Tepals 4–5 by 0.6–1.2 mm. Column straight to strongly curved at base,

cylindrical, without wings. Stamens 2.3–4 mm long. Staminode entirely adnate to column

or with free obtuse tip. Ovary 1–1.7 cm long, 0.8–1 mm in diam. Capsule 1–1.8 cm

long, 1.4–2 mm wide. Fig. 15, a, b.

Ecology. Shady broad-leaved evergreen and coniferous lowland and submontane

forests on shale, sandstone, granite, gneiss and basalt. 200–1200 m. Fl. March,

April. Occasional (LR).

Distribution. Vietnam (Dong Nai, Gia Lai, Khanh Hoa, Kon Tum, Lam Dong,

Ninh Thuan, Phu Tho, Quang Binh, Quang Nam, Tay Ninh, Thua Thien – Hue).

Tropical Asia from Sri Lanka and Nepal to Philippines, N Australia and New Guinea.

Studied specimens. Prenh, Dalat, Evrard, 1469, 2314 (P); Lo Go, Tay Ninh,

Mueller, 1470 (P); Gian Lau me, Dong Nai, Bienhoa, Pierre, 6583 (P); Mt Tamire, Pierre, 6583

(P); Nhatrang, Poilane, 3808 (P); Ba Na, Danang, Poilane, 7277 (P); Go O to Mo O, Quang

Nam 1000 m, Poilane, 31671 (P); Gialai – Kontum, Kbang – Kon Ha Nung, LX-VN 2739 (LE);

Khanh Hoa, Khanh Son, Hon Giao mt., VH 4322 (HN, LE); Kon Tum, Kon Plong, DKH 4690

(HN, LE); Kontum, Kon Plong, Mang La, VH 5185 (HN, LE); Kontum, Kon Plong, Mang La, VH

5327 (HN, LE); Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4321 (HN, LE), HLF 4362 (HN, LE);

Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4362 (HN, LE); Thua Thien – Hue, A Luoi, HAL 7968

(HN, LE); Quang Binh, Minh Hoa, HAL 11708 (HN).

2. A. odorata Blume, 1825, Bijdr. 6, fig. 39; id. 1825, ibid. 8: 423; Seidenf.

1992, Orch. Indochina: 13; Aver. 1994, Identif. Guide Viet. Orch.: 13; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 11. – A. thorelii Gagnep. 1933,

Bull. Soc. Bot. France, 70: 350.

Described from Java (“montis Salak”). Lectotype (“Blume s.n.”) – L.

Plant 30–80 cm tall. Leaves numerous, close, narrowly lanceolate to lanceolate,

15–25 by 1.5–2.2 cm. Inflorescence simple to few branched, many flowered. Flowers

bright yellow. Tepals 9–12 by 1.5–2 mm. Column straight to strongly curved at base,

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 13. Neuwiedia zollingeri var. balansae: a – flower. Apostasia odorata: b – flowering

plant, c – flower, d – column; A. nipponica: e – flowering and fruiting plant, f – flower. A. nuda:

g – flowering plant.


Turczaninowia 2008, 11(1) : 5–168

Fig. 14. Neuwiedia zollingeri var. javanica: a, b (HLF 4415), c, d (HLF 5479). N. zollingeri

var. balansae: e, f (HAL 1686), g (HLF 5567).

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 15. Apostasia wallichii: a, b (VH 5185). A. odorata: c-e (HAL 5337). A. nipponica: f (HLF

1406), g, h (HAL 7175).


Turczaninowia 2008, 11(1) : 5–168

usually laterally flattened, with 2 short longitudinal low to tall quadrangular wings below

staminode. Stamens 2, 5–6 mm long. Staminode adnate to column with free obtuse

tip. Ovary 10–18 mm long, 1–1.2 mm wide. Capsule 1.8–2.5 cm long, 1.5–2.5 mm

wide. Fig. 13, b-d; 15, c-e.

Ecology. Shady broad-leaved evergreen and coniferous lowland, submontane

and montane forests on shale, sandstone, granite, gneiss, rhyolite and quartzite. 300–

1600 m. Fl. March – June. Occasional (LR).

Distribution. Vietnam (Dak Lak, Dong Nai, Ha Tinh, Khanh Hoa, Lam Dong,

Lao Cai, Phu Tho, Quang Binh, Vinh Phuc). Mainland tropical Asia, Hainan, Java,

Sumatra, Kalimantan.

Studied specimens. Bao Loc 1000–1200, Poilane, 22045 (P); Bavi, 1200 m,

Chevalier, 37777 (P); Dak Lak, Chu Yang Sin mt., HLF 5481 (HN, LE); Ha Tinh, Huong Son,

HAL 1281 (HN, LE), HAL 5337 (HN, LE); Khanh Hoa, Khanh Son, Hon Giao mt., VH 4174 (HN,

LE), VH 4378 (HN, LE); Lao Cai, Van Ban, DKH 6626 (HN, LE), DKH 6905 (HN, LE); Lao Cai,

Van Ban, HAL 2354 (HN, LE); Vinh Phu, Tam Dao 3, LX-VN 2368 (LE); Quang Binh, Quang

Ninh distr., HAL 11519 (HN); Quang Binh, Minh Hoa, HAL 11624 (HN).

Notes. The name A. thorelii is attributed to plants with tall quadrangular

column wings and rather broad leaves. Taxonomical status of such marginal plants is

still uncertain.

3. A. nipponica Masam. 1935, Journ. Jap. Bot. 11: 46.

Described from Ryukyu Islands (“Yakusima, Kurio, …”). Type (“Jul. 13. 1933

(leg. S. Suzuki)”) – ?

Plant 5–10(12) cm tall. Leaves few, distant, distinctly petiolate, petiole 0.5–1

cm long, leaf blade ovate to elliptic, 1.5–2.5 by 0.5–1 cm. Inflorescence simple or with

1–3 branches bearing 1–4 flowers. Flowers bright yellow. Tepals 3.5–5.5 by 1–1.4

mm. Column straight to slightly curved at base, without wings or with 2 short longitudinal

insignificant wings below staminode. Stamens 2.2–3 mm long. Staminode adnate to

column with free orbicular tip. Ovary 6–9 mm long, 0.5–1.2 mm wide. Capsule 0.8–

1.1 cm long, 1.4–1.8 mm wide. Fig. 13, e, f; 15, f-h.

Ecology. Shady broad-leaved evergreen lowland and submontane forests on

shale, sandstone and granite. 300–1000 m. Fl. April, May. Very rare (LR).

Distribution. Vietnam (Ha Tinh, Thua Thien – Hue). Ryukyu Islands.

Studied specimens. Ha Tinh, Huong Son, Rao An, No HAL 5157 (HN, LE); Thua

Thien – Hue, A Luoi, HAL 7175 (HN, LE); Thua Thien Hue, Bach Ma, HLF 1406 (HN, LE);

Thua Thien Hue, Nam Dong, HAL 10928 (HN, LE).

4. A. nuda R. Br. 1830, in Wall. Pl. As. Rar. 1: 76, tab. 85; Seidenf. 1992,

Orch. Indochina: 15; Aver. 1994, Identif. Guide Viet. Orch.: 13; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 11.

Described from Malacca Peninsular (“in montibus insulae Penang”).

Lectotype (“Porter s.n. Wallich 4449”) – K.

Plant 12–40 cm tall. Leaves numerous, close, linear to linear lanceolate, 7–20

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

by 3–8 mm. Inflorescence simple to few branched, many flowered with tuft of small

narrow sterile bracts at base. Flowers white to yellow. Tepals 3.5–4.5 by 0.6–0.8 mm.

Column strongly curved. Stamens 2, 3.2–3.5 mm long. Staminode absent. Ovary 8–

10 mm long, 0.8–1 mm wide. Capsule 1.2–1.4 cm long, 1 mm wide. Fig. 13, g.

Ecology. Shady coniferous and broad-leaved evergreen lowland and mountain

forests on shale, 320–1400 m. Fl. January, February. Very rare (DD).

Distribution. Vietnam (Dak Lak, Thua Thien – Hue). Myanmar, Thailand,

Malacca Peninsular, Java, Sumatra, Kalimantan.

Studied specimens. Dak Lak, Chu Yang Sin mt., HLF 5438 (HN, LE); Thua Thien –

Hue, Nam Dong, HAL 6983 (HN, LE).

Notes. Flowers of this species varies from yellow to white. Flower color of

Vietnamese samples is unknown.

SUBFAMILY CYPRIPEDIOIDEAE LINDL.

1821, Collect. Bot. Cypripedieae (sub “Cypripedieae”). - Fam. Cypripediaceae

Lindl., 1833, Nixus Pl.: 22 (sub “Cypripedieae”).

Type: Cypripedium L.

5(1) genera and about 125(18) species. Subtropical and tropical areas of the

world (except Africa).

Paphiopedilum Pfitz.1886, Morph. Stud. Orch.: 11; Stein, 1892, Orchideenbuch:

448–492; Seidenf. 1972, Bot. Tidsskr. 67, 1–2: 76–95; Chen et Liu, 1982, Acta Bot.

Yunnan. 4, 2: 163–167; Braem, 1988, Paphiopedilum: 1–249, Cribb, 1998, Gen.

Paphiopedilum: 1–427; Averyanov et al. 2003, Slipper Orch. Vietnam: 1–288, nom. cons.

Type: P. insigne (Wall. ex Lindl.) Pfitz. (Cypripedium insigne Wall. ex Lindl.)

Sympodial terrestrial, lithophytic or epiphytic herbs with thick hairy roots at

the base of very short stem. Leaves few, rosulate, distichous, conduplicate, often

tessellated with deep and light green mottles. Inflorescence sub-terminal, one- to fewflowered

raceme on long stalk. Flowers large, resupinate, variously colored, pubescent

outside. Lateral sepals connate into synsepalum. Lip inflated, sac-like, slipper-shaped.

Column short, curved downwards into lip opening, dorsally with large shield-like

staminode, 2 lateral fertile stamens and hemispheric stigma ending short style. Pollen

not aggregated into pollinia, gel-like, sticky.

About 75(18) species. Tropical Asia to Philippines and islands of west Pacific.

Key to species

1. Lip inflated, sub-spherical to ovoid, calceolate with involuted upper apical margin;

dorsal sepal and synsepal more or less similar to petals in color and shape, white,

yellowish, dull yellow-greenish or pink, more or less of one color; petals broadlyovate

to circular, less than twice as long as broad, very rarely petals narrowly-ovate

(in this case petals and lip hooked at apex) ....................................................................... 2

– Lip goblet-like, cylindrical, with incurved side-lobes and apical margin of the pouch


Turczaninowia 2008, 11(1) : 5–168

straight, not involuted; dorsal sepal and synsepal distinctly differ from petals in color

and shape, petals much narrower than dorsal sepal, dorsal sepal and petals commonly

variously colored, often contrastly striped and spotted; petals tapering, narrowlyoblong

or spathulate, more than twice as long as broad, newer hooked at the apex ......... 8

2. Dorsal sepal and synsepal distinctly smaller than lateral petals; lip inflated, more or

less spherical, thin-textured, lightly grooved along the veins; staminode large, broad

at apex, flat or longitudinally conduplicate, sometimes with involute side margins;

flowers white, pale yellowish-green or pink, sometimes with brown or purple stripes

along nerves; leaves tessellated or uniformly green; plant of calcareous or acidic

silicate substrates ................................................................................................................ 3

– Dorsal sepal and synsepal more or less similar with petals size; lip is ovoid, rather

thick in texture, plain on surface; staminode middle sized, distinctly 3- or 5-dentate at

the apex, with a large narrow central tooth; flowers pale yellowish-white to yellow,

sepals and petals usually with small purple spots; leaves markedly tessellated; plant

of calcareous substrates ...................................................................................... 7. P. concolor

3. Leaves distinctly tessellated above and heavily spotted or uniformly tinged with

dirty-purple-violet beneath, 10–15(20) cm long, 1.5–4 cm wide; flowers pink, pinkpurple

or pale green; lip longer or hardly shorter than lateral petals; underground

stolons may be present or not; plants of calcareous or acidic silicate substrates .... 4

– Leaves uniformly green on both sides, rare with few small purple-violet marks beneath

near the base, (15)20–35 cm long, 4–6 cm wide; flowers white or pale yellowish-green;

lip distinctly smaller than lateral petals; plants of calcareous substrates, never

developing stolons .............................................................................................................. 7

4. Staminode flat, with a broad apex; sepals and petals uniformly light pink to pinkpurple;

plants of calcareous or acidic silicate substrates, without stolons .................. 5

– Staminode longitudinally conduplicate or with deep fold along midvein at the apex;

sepals apple-green or yellowish-pink with purple stripes along veins; plants of

calcareous substrates, with more or less elongate underground stolons ..................... 6

5. Flowers 7.5–10 cm across; sepals less than 4.5 cm long, subcircular to broadly-ovate,

spreading; staminode with bright yellow center and central purple mark; leaves

narrowly-oblong to oblong, 8–11 cm long, 3–3.5(4) cm wide, tessellated more or less

contrastingly deep-green/whitish-green; plant of acidic silicate substrates ..... 1. P. delenatii

– Flowers 10–12 cm across; sepals longer than 4.5 cm, broadly-oblong to broadlyovate,

down directed; staminode with dull yellow or light olive-green center; leaves

ovate to oblong-elliptic, (8)11–20 cm long, (2.5)4–7 cm wide, tessellated glossy deepgreen/silver-gray;

plant of calcareous substrates ..................................... 2. P. vietnamense

6. Flowers (8)9–12.2 cm across, pale yellowish-green to apple-green, dorsal sepal and

petals with brown-purple stripes; dorsal sepal cuspidate-acuminate at apex, more

than 4 cm long; petals ovate to narrowly-ovate, subacute at apex, longer than 4 cm;

staminode more than 12.5 mm long, broadly ovate, convex at the base, apically with a

deep fold along the midvein and involute side margins ............................. 3. P. malipoense

– Flowers 6–8(8.7) cm across, pink to pink-purple dorsal sepal and petals with purple

striped along veins; dorsal sepal obtuse or shortly acute at apex, less than 4 cm long;

petals broadly obovate to subcircular, rounded at apex, less than 4 cm long; staminode

less than 12.5 mm long, elliptic, longitudinally conduplicate ................... 4. P. micranthum

7(3). Flowers commonly subcampanulate, sepals and petals light yellowish-green with

purple-brown flushing at the base; lip yellowish-white; staminode broadly triangular,

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

more than 17 mm wide, convex, broadly-rounded at apex, white or yellowish-white

with purple anastomosing markings ............................................................. 5. P. hangianum

– Flower commonly spreading, sepals and petals white, sometimes faintly flushed with

purple at base; lip yellow to pink-orange, with a pink flush around the rim; staminode

narrowly-triangular, less than 12 mm wide, longitudinally sulcate, with a deep groove

along midvein and narrow apex, bright yellow with reddish markings and a white

margin ............................................................................................................. 6. P. emersonii

8(1). Inflorescence one-flowered, very rarely with 2 flowers; petals narrowly-obovate to

oblong-spathulate, more or less flat, undulate at margin but never spirally twisted,

less than 8(8.5) cm long; medium-sized or small plants with more or less herbaceous

leaves commonly less than 20 cm long, very rarely longer ............................................. 9

– Inflorescence with (1)2–5 flowers; petals linear-tapering, strongly spirally twisted,

much narrower than median sepal, 8–11 cm long; large plants with coriaceous leaves

20–50 cm long, 2–5 cm wide ............................................................................. 18. P. dianthum

9. Leaves uniformly green, sometimes with deep green venation; often with purpleviolet

marks beneath near the base; staminode obovate, obcordate, subcircular or

subquadrate with a central umbo or broad convex callus; petals without maroonviolet

warts; lip with plain smooth surface of side-lobes (P. sect. Paphiopedilum) ... 10

– Leaves distinctly tessellated deep-green/light-green, without purple-violet marks

beneath; staminode commonly lunate, with falcate side-lobes and a small central

tooth, rare broadly obcordate, 3- or 5-dentate at the apex, without distinct central

boss; petals with more or less prominent small maroon-violet warts along margin and

near the base; lip with warted surface of side-lobes ...................................................... 16

10. Staminode subquadrate, smooth, glossy, with a broad central swelling, lacking a glossy

umbo; petals purple in apical half ............................................................ 8. P. hirsutissimum

– Staminode obovate, obcordate or semicircular, rugulose, with a central glossy umbo;

petals orange, yellow, yellow with a pink tint, yellowish-brown or light brown .......... 11

11. Flowers small or medium-sized, less than 6.5 cm across; dorsal sepal shorter than

4cm; petals less than 4 cm long ....................................................................................... 12

– Flowers large, more than (7) 7.5 cm across; dorsal sepal longer than 4.5 cm; petals

more than 4.5 cm long ........................................................................................................ 15

12. Plants small with coriaceous, rigid, semi-succulent leaves 3–7 (12) cm long; flowers

bright yellow to dull yellowish-pink with an ochre-yellow to dull yellowish-pink lip;

petals with straight margins, commonly ochre-yellow with orange veins ... 12. P. helenae

– Plants medium-sized with more or less herbaceous leaves more than 7 cm long; flowers

not colored as above; petals with more or less undulate margins, light yellowishbrown

or dull yellowish with a pink tint or brown veins ................................................ 13

13. Petals oblong, with strongly undulate margins, light yellowish-green with chocolatebrown

veins and flushing; lip chocolate-brown with darker veins; dorsal sepal white

with a greenish base and chocolate-brown veins towards the base .... 10. P. tranlienianum

– Petals spathulate or narrowly obovate, with slightly undulate margins, light yellow-brown or

dull yellowish with pink tint and large maroon spots; lip pink-purple or purple-brown; dorsal

sepal white with a purple center or dull yellowish with maroon spots .................................. 14

14. Dorsal sepal subcircular to obovate, with reflexed margins at the base, white with a

purple or brownish-purple center; petals narrowly-spathulate, 0.8–1 cm wide, dull

yellowish-brown with pink-purple tint; lip yellowish-brown to purple-brown ....................

............................................................................................................................ 9. P. barbigerum


Turczaninowia 2008, 11(1) : 5–168

– Dorsal sepal broadly-ovate to ovate, more or less flat, with flat margins at the base,

light dull yellowish-green, heavily spotted with large maroon spots; petals broadlyoblong

to narrowly-obovate, 1.4–1.6 cm wide, dull yellowish, flushed with pink-purple,

spotted with numerous large maroon spots; lip pink-purple ................... 12. P. henryanum

15(11). Dorsal sepal 4.8–5.2 cm long, white, sometimes with light brownish or green at the

base, more or less heavily spotted with brown-purple; petals spathulate, usually less

than 5.5 cm long, 1.8–2.5 cm wide; peduncle commonly erect, shortly densely hairy

with short purple-violet hairs; lithophytic plant .................................... 13. P. gratrixianum

– Dorsal sepal (4.5) 5–7 cm long, greenish with a white margin and glossy maroon

central area, rare maroon spotted; petals obovate-spathulate, usually more than 5.5cm

long, 2.5–4.6 cm wide; peduncle commonly arching, densely villosely hairy with long

whitish hairs; epiphytic plant, very rare a lithophyte .................................... 14. P. villosum

16(9). Dorsal sepal pale green with darker venation, forming a hood arching over the lip;

staminode broadly cordate with 1–3 obtuse apical teeth or lunate with broad falcate

side-lobes, 3- or 5-denticulate at the apex; petals spathulate, broadening toward apex;

lip often with 2–6 obtuse teeth on front margin ................................. 15. P. appletonianum

– Dorsal sepal erect, more or less flat, with reflexed margins at the base, white with

purple or green veins; staminode lunate with narrow falcate side-lobes and 1–3 small

median teeth; petals oblong, sigmoid or elliptic; lip front margin without obtuse teeth ... 17

17. Petals more or less sigmoid, down-curved, green with a pink-purple apex and prominent

deep maroon hairy warts along the upper margin; dorsal sepal subcircular to broadlyelliptic,

white, green/purple striped; lip pink-brown to dull purple; leaves commonly

elliptic to obovate, 10–20 cm long ................................................................... 16. P. callosum

– Petals elliptic, horizontally spreading, deep-purple, finely purple-violet spotted in

lower part, without hairy warts along the upper margin; dorsal sepal broadly-ovate,

white, deep-purple striped; lip brown-purple to deep purple-violet; leaves commonly

oblong to elliptic, 7–14 cm long ................................................................... 17. P. purpuratum

1. P. delenatii Guillaum. 1924, Bull. Soc. Bot. France, 71: 554, 558; Seidenf.

1992, Orch. Indochina: 16; Aver. 1994, Identif. Guide Vietnam. Orch.: 16; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003, Slipper Orch.

Vietnam: 89. – Cypripedium delenatii (Guillaum.) C.H. Curtis, 1931, Gard. Chron.

Ser. 3, 89: 208.

Described from Vietnam (“Tonkin: Sans localite precise. ...; Annam: Nha trang”).

Lectotype (“Tonkin, Delenat, cult. Mornay, Fleuriste de la ville de Paris”) – P.

Lithophytic or terrestrial herb with 5–7 oblong-elliptic leaves up to 11 by 3–3.9

cm, distinctly tessellated, heavily purple-spotted below. Peduncle with 1–2(3) flowers,

purple-spotted, up to 22 cm long; pedicel and ovary up to 5.5 cm long, spotted with

purple, white hairy. Flower 7.5–8 cm across, pale pink with pink or purple-pink lip.

Median sepal ovate, 1.7–3.5 by 1.8–2.5 cm. Synsepal similar, 1.9–3 by 1.4–2.9 cm.

Petals broadly elliptic, obtuse, 3–4.2 by 2.4–3.8 cm. Lip ellipsoidal to subglobose, 2.5–

3.8 by 2.5–3 cm. Staminode convex, ovate, 14–17 by 13–16 mm with red and yellow

markings. Fig. 16, a-c; 23, a.

Ecology. Shady, broad-leaved, evergreen lowland and submontane forests on

granite and gneiss 750–1300 m. Fl. November, December. Very rare (CR).

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Distribution. Vietnam (Dac Lac, Khanh Hoa). Endemic.

Studied specimens. Khanh Hoa, Lieng Ly River, L. Averyanov et al. s.n., Apr. 1995

and Dec. 1995 (HN); Khanh Hoa, Khanh Son, Hon Giao mt., VH 4304 (HN); Lam Dong, Dalat

area, Averyanov s.n., 2005 (LE – photo).

Notes. Local endemic with range area less than 100 square km., closely

allied to calcium dependent Paphiopedilum armeniacum and P. vietnamense. Flowers

commonly vary from light pink to pink with purple lip. Individual plants in some

populations have white or nearly white flowers.

2. P. vietnamense Gruss et Perner, 1999 (11 Jan.), Die Orchidee Beihefte, 5:

3; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 49; Aver et al. 2003,

Slipper Orch. Vietnam: 96. – P. hilmari Senghas et Schettler, 1999 (19 Jan.), Journ.

Orchideenfr. 6, 1: 4. – P. mirabile Cavestro et Chiron, 1999 (19 Feb.), Orch. Cult.

Protect., 32, 2: 32.

Described from northern Vietnam, Thai Nguyen province (“Cao Bang Province,

1000 m” – exactly wrong). Type (“fl. in cult. November 1998, Dec. 1998, O. Gruss”) –K.

Lithophytic or terrestrial herb with 3–5 glossy, oblong-elliptic leaves 8–17(20)

by 2.5–7 cm, distinctly tessellated, spotted or flushed with purple beneath. Peduncle

erect, 1(2)-flowered, 15–25 cm long, purple-violet to purple-brown, white hairy. Flower

10–12 cm wide, pink to purple; lip commonly deep purple on front or around rim.

Pedicel and ovary 2.5–2.8 cm long, fine purple spotted. Median sepal erect, ellipticobovate,

4.2–5.1 by 2.5–3 cm. Synsepal elliptic-ovate, 3–4.6 by 3–3.3 cm. Petals

deflexed or spreading, oblong-elliptic, 5–6 by 3.3–4.5 cm. Lip subglobose, 3.5–4 cm

across. Staminode large, convex, broadly rhombic, lemon-yellow with white margin

and olive-green center. Fig. 16, d, e; 23, b.

Ecology. Shady broad-leaved evergreen humid forests on rocky crystalline

limestone 350–550 m. Fl. March. Extinct in nature (EW).

Distribution. Vietnam (Thai Nguyen). Endemic.

Studied specimens. Thai Nguyen, Dong Hy, HAL 31 (HN).

Notes. Bright example of restricted local endemic completely depleted in

nature by commercial collectors. All known specimens originate from alone miserable

area in Thai Nguyen province. The species presently known only in cultivation is quite

variable in both coloration and shape of its flowers.

3. P. malipoense S.C. Chen et Z.H. Tsi, 1984, Acta Phytotax. Sin. 22, 2: 119;

Aver. 1994, Identif. Guide Vietnam. Orch.: 16; Aver. et Averyanova, 2003, Updated

Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam: 104.

Described from S China (“Yunnan: Malipo, Hwang Jin In, 1300–1600 m”).

Type (“Nov. 11 1947, K.M. Feng 13161”) – PE.

Distribution. N Vietnam, S China, NE Laos.

Key to varieties

1. Petals broadly obovate to obovate; lip wider than petals, 3.8–5.4 cm wide; staminode


Turczaninowia 2008, 11(1) : 5–168

with deep brown-purple or maroon apex; leaves heavily spotted with purple-violet

beneath .............................................................................. 3a. P. malipoense var. malipoense

– Petals obovate to narrowly obovate or lanceolate; lip of equal width or narrower than

petals, usually less than 3.8 cm wide; staminode with white, light yellowish or light

greenish apex marked with finest brown-purple marks; leaves sparsely spotted with

purple-violet beneath .......................................................................................................... 2

2. Petals obovate to narrowly obovate; lip subglobose, more than 1.8 cm wide, truncate

at the apex .................................................................................. 3b. P. malipoense var. jackii

– Petals narrowly obovate to lanceolate; lip narrowly ovate, slightly bilaterally

compressed, hooked at apex ................................................... 3c. P. malipoense var. hiepii

3a. P. malipoense var. malipoense.

Lithophytic or terrestrial herb with creeping rhizome and 4–6 coriaceous oblong

leaves 10–16(20) by 2.5–5(7) cm, distinctly tessellated, heavy purple-marked below.

Peduncle 30–65 cm tall, 1(2)-flowered, purple-brown spotted, white hairy. Flower 8–

12.2 cm across, apple fragrant, light green with brown striped tepals. Pedicel and

ovary about 4 cm long, white hairy. Tepals brown striped. Dorsal sepal ovate, 4.4–7.1

by 1.8–4.5 cm. Synsepal ovate, 3.8–5.3 by 2.4–4.8 cm. Petals ovate, 4–7.1 by 3.4–

5.1 cm. Lip pale yellow-green to greyish, subglobose, 4–6.5 cm across. Staminode

convex, ovate-oblong, 1.3–1.4 by 1.1–1.3 cm wide, pure white, with truncate deep

maroon-brown apex. Fig. 16, f-h; 23, c.

Ecology. Shady broad-leaved evergreen and coniferous humid forests on rocky

crystalline limestone. 400–1450 m. Fl. March, April. Occasional (EN).

Distribution. Vietnam (Bac Kan, Ha Giang, Hoa Binh, Lang Son, Quang

Binh, Son La, Thanh Hoa, Tuyen Quang). S China, NE Laos.

Studied specimens. Bac Kan, Na Ri, NTH 3705 (LE), DKH 7554 (HN, LE, MO);

Bac Kan, Cho Don, HAL 4857 (HN, LE); Ha Giang, Meo Vac, HAL 8498 (HN); Ha Giang, Quan

Ba, HG 52 (HN, LE), NTH 3580 (HN, LE), DKH 4805 (HN, LE), HAL 8338 (HN), HAL 8377

(HN); Hoa Binh, Da Bac, HAL 350 (HN, LE); Hoa Binh, Mai Chau, HAL 703 (HN, LE), HAL 832

(HN, LE), HAL 871 (HN, LE), N.T. Hiep et al., s.n. (HN, LE), NTH 2040 (HN, LE), VH 2413 (HN,

LE), DKH 7707 (HN, LE, MO), DKH 7992 (HN, LE, MO), DKH 7799 (HN, LE, MO), DKH 8058

(HN, LE, MO); Lang Son, Huu Lung, NTH 3258 (HN, LE); Quang Binh, Bo Trach, HAL 6117

(HN), HAL 6207 (HN); Quang Binh, Minh Hoa, VH 4759 (HN, LE), HAL 5839 (HN), HAL 5967

(HN, LE), HAL 6039 (HN); Son La, Moc Chau, DKH 7418 (HN, LE), HAL 9296 (HN, LE); Son

La, Yen Chau, HAL 9434 (HN, LE), HAL 9505 (HN); Thanh Hoa, Ba Thuoc, HAL 974 (HN, LE),

HAL 1047 (HN, LE), HAL 3287 (HN), HAL 3438 (HN); Thanh Hoa, Quan Hoa, HAL 3934 (HN),

HAL 3935 (HN); Tuyen Quang, Na Hang, HAL 245 (HN); Quang Binh, Minh Hoa, HAL 11688

(HN), HAL 11735 (HN).

Notes. Very sensitive species, which dies out very fast under deforestation. It

is quite variable both in coloration and in shape of its flowers. Albino forms and forms

with more deep coloration time to time are recognizing in imported materials.

3b. P. malipoense var. jackii (H.S. Hua) Aver. 1997, Orchids, 66, 2: 153. –

P. jackii H.S. Hua, 1996, Die Orchidee, 46, 3: U4.

Described from S China (“Yunnan, Wenshan”) – questionable. Type (“12

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 16. Paphiopedilum delenatii: a – flowering plant and flower, b, c – staminode; P. vietnamense:

d – flowering plant, e – staminode. P. malipoense var. malipoense: f – flowering

plant, g, h – staminode (HAL 6207). P. malipoense var. hiepii: i – flower, j – lip, k, l – staminode.


Turczaninowia 2008, 11(1) : 5–168

March 1995, S.S. Hu 3268 A-S”) – IBSC.

Lithophytic or terrestrial herb. Leaves light green, lightly tessellated and veined

dark green above, light green and sparsely spotted with purple-violet beneath. Flowers

differ from the typical variety in having narrower petals, a smaller yellowish-green or

green lip and a white staminode, usually with a yellow, yellow-green or green central

mark with fine purple venation. Fig. 23, d.

Ecology. Broad-leaved evergreen dry, rather open forests on stratified soft

limestone. 550–650 m. Fl. March, April. Very rare (EN).

Distribution. Vietnam (Tuyen Quang), S China?

Studied specimens. Tuyen Quang, N.T. Hiep s.n. 1995 (LE); Tuyen Quang, Na

Hang, HAL 169 (HN, LE), HLF 661 (HN, LE).

Notes. Flowers are remarkably variable in the form of petals and coloration

of staminode, which may be pure white, yellowish or yellow-green with fine purple

venation and marks. In fact, the variety is credibly known only from small area in

northern Vietnam in Tuyen Quang province. Reports of this species from China may

be based on imported plants originated from this area.

3c. P. malipoense var. hiepii (Aver.) P.J. Cribb, 1998, Genus Paphiopedilum:

88. – P. hiepii Aver. 1998, Orchids, 67, 3: 261. – P. jackii H.S. Hua var. hiepii (Aver.)

Koop. 2000, Orchid Digest, 64, 4: 168. – P. angustatum Z.J. Liu et S.C. Chen, 2000,

Acta Phytotax. Sin. 38, 5: 464.

Described from N. Vietnam (“Prov. Tuyen Quang, Distr. Tuyen Quang”).

Type (“14–15 June 1995, N.T. Hiep s.n.”) – LE.

Terrestrial herb. Leaves nearly white with dark green venation above and

sparse purple-violet spotting beneath. Flowers differ from the typical variety in narrow

ovate or lanceolate petals hooked at the apex, a small narrow laterally compressed

yellowish-green lip and a white staminode with few fine purple marks at apex. Fig. 16,

i-l; 23, e.

Ecology. Shady broad-leaved evergreen dry forests on soft slate limestone

and schist. 500–600 m. Fl. March, April. Extinct in nature (EW).

Distribution. Vietnam (Son La?, Tuyen Quang). S China?

Notes. Species is known only by type collection. Report from S China may be

based on imported plants originated from locus classicus area. Doubtful plants with

similar coloration of leaves were occasionally observed in Son La province (HAL 9555).

4. P. micranthum T. Tang et F.T. Wang, 1951, Acta Phytotax. Sin. 1, 1: 25,

56; Aver. 1994, Identif. Guide Vietnam. Orch.: 17; Aver. et Averyanova, 2003, Updated

Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam: 120.

Described from S China (“Yunnan”). Type (“C.W. Wang 86182”) – PE.

Lithophytic or terrestrial herb with creeping rhizome and 3–5 oblong rigid leaves

5–12(15) by 1.5–2 cm, distinctly tessellated, purple-violet spotted below. Peduncle

erect, 1-flowered, 9–25 cm long, purple-brown spotted, white hairy. Flower odorless,

5.8–9 cm tall, thin-textured; tepals yellowish flushed with pink, veined with deep red-

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purple. Median sepal ovate, 1.5–3.6 by 1.7–3 cm. Synsepal elliptic, 1.4–3.3 by 1.1–

2.5 cm. Petals broadly obovate, 1.9–4.3 by 2.3–4.4 cm, ciliate with long white hairs.

Lip white to pink-purple, deeply inflated, 5–10.4 by 3.4–5.6 cm. Staminode white,

yellow above, spotted with red, conduplicate, 10–12.5 mm long. Pedicel and ovary

3.5–4.6 cm long, brown-purple spotted, white hairy. Fig. 17, a-d; 23, f.

Ecology. Evergreen and semi-deciduous, mixed and coniferous forest on rocky

limestone. 600–1600 m. Fl. March, April. Rare (EN).

Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Tuyen Quang), S China.

Studied specimens. Bac Kan, Cho Don, HAL 4771 (HN, LE), HAL 4794 (HN, LE),

HAL 4864 (HN, LE), HLF 796 (HN, LE); Cao Bang, Bao Lac, CBL 1477 (HN, LE), CBL 332

(HN); Cao Bang, Ha Lang, CBL 627 (HN, LE), CBL 767 (HN, LE); Cao Bang, Nguyen Binh,

CBL 555 (HN, LE), CBL 1277 (HN, LE), CBL 1577 (HN, LE); Cao Bang, Tra Linh, CBL 1079

(HN, LE), CBL 1222/2 (LE); Cao Bang, Trung Khanh, HAL 5457 (HN), HAL 5643 (HN, LE),

HAL 5685 (HN), HAL 5745 (HN); Ha Giang, Dong Van, CBL 1776 (HN, LE), NTH 3346 (HN,

LE), HAL 8594 (HN), HAL 8615 (HN), HAL 8653 (HN), HAL 8689 (HN); Ha Giang, Meo Vac,

CBL 1822 (HN, LE), NTH 3432 (HN, LE), HAL 8504 (HN), HAL 8564 (HN); Ha Giang, Quan

Ba, NTH 3576 (HN, K, LE), DKH 6173 (HN, LE), HAL 8341 (HN); Ha Giang, Vi Xuyen, DKH

6451 (HN, LE); Ha Giang, Yen Minh, CBL 1940 (HN, LE), NTH 3494 (HN, LE); Tuyen Quang,

Na Hang, HAL 147 (HN), HLF 659 (HN, LE).

Notes. Very sensitive species, which vanish very fast under deforestation. It

is quite variable in coloration of leaves and flowers. Its populations occasionally include

albino forms and forms with more deep flower coloration.

5. P. hangianum Perner et Gruss, 1999, Die Orchidee, Biehefte, 6: 3; Aver.

et Averyanova, 2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper

Orch. Vietnam: 131. – P. singchii Z.J. Liu et J.Y. Zhang, 2001, Acta Phytotax. Sin.

38, 5: 468.

Described from N Vietnam (“Bac Thai Province, Tong Ngoc Hang s.n.”).

Type (“cult. Gruss HAL 072655”) – HAL.

Lithophytic herb with 4–7 coriaceous, ligulate, glossy, green leaves (12)15–30

by 3–5 cm. Peduncle erect, 1(2)-flowered, 6–12 cm long, shortly white hairy. Pedicel

and ovary 3.5–4.2 cm long, bright green. Flower slightly sweetly fragrant, 9–12 cm

wide, rather thick-textured, light yellowish or pale yellowish-green. Median sepal ovate,

4–5.6 by 2.2–3.9 cm. Synsepal broadly ovate, 3.5–6 by 3–4.7 cm. Petals broadly

elliptic, 4–7 by 2.5–4.5 cm. Lip yellowish-white, ovoid, 3.3–5 by 2.3–3.4 cm. Staminode

broadly sagittate, white or yellowish-white with purple anastomosing markings, 1.1–

1.7 by 1.1–2.1 cm wide. Fig. 17, e, f; 23, g.

Ecology. Shady broad-leaved evergreen dry forests on soft stratified limestone.

450–750 m. Fl. May, June. Rare (EN).

Distribution. Vietnam (Bac Kan, Tuyen Quang). Endemic.

Studied specimens. Bac Kan, Cho Don, HAL 4789 (HN), HLF 806 (HN, LE);

Tuyen Quang, Na Hang, HAL 126 (HN, LE), HAL 192 (HN, LE).

Notes. Local endemic with very restricted distribution. There are few doubts


Turczaninowia 2008, 11(1) : 5–168

that description of P. singchii reported from southern Yunnan is based on plants imported

from Vietnam.

6. P. emersonii Koop. et P.J. Cribb, 1986, Orchid Advocate, 12: 86; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003, Slipper Orch.

Vietnam: 140. – P. huonglanae N.T. Tich, 1998, Hoa Canh (1998), 3: 10–11, nom.

inval. – P. emersonii Koop. et P.J. Cribb var. huonglanae (N.T. Tich) N.T. Tich,

1998, Hoa Canh (1998), 4: 12, comb. inval.

Described from China (“China”). Type (“cult. E.W. Charles”) – K.

Lithophytic herb with 4–7 coriaceous, ligulate, glossy green leaves, below with

sparse purple-violet marks near the base. Peduncle erect, 1-flowered, 10–15 cm long,

yellowish-green, shortly white hairy. Pedicel and ovary about 3 cm long. Flower slightly

sweet fragrant, 8–10.5 cm wide, rather thick-textured. Median sepal ovate, 2.7–5.1

by 1.5–3.5 cm. Synsepal broadly elliptic, 2–4.5 by 1.8–4 cm. Petals broadly obovate,

3.5–5.4 by 2–4 cm, white villose and sometimes faintly flushed with pink at base. Lip

light yellow to dull orange-pink, obovoid, 3–5 by 2–3 cm. Staminode bright yellow with

reddish markings, 1.6–2 by 0.8–1.1 cm, deeply longitudinally sulcate. Fig. 17, g-i; 23, h.

Ecology. Shady broad-leaved evergreen dry forests on rocky solid slaty limestone.

550–750 m. Fl. May, June. Very rare (CR).

Distribution. Vietnam (Bac Kan, Thai Nguyen, Tuyen Quang). S China.

Studied specimens. Bac Kan, Na Ri, HAL 4963 (HN), DKH 7644 (HN, LE, MO);

Thai Nguyen, Dong Hy, HAL 031/1 (LE); Thai Nguyen, Vo Nhai, HAL 1581 (HN).

Notes. Variable species, particularly in size of flowers, petal shape and color

of the lip.

7. P. concolor (Bateman) Pfitz. 1889, in Engler et Prantl, Natur. Pflanzenf. 2,

6: 84; Seidenf. 1992, Orch. Indochina: 16; Aver. 1994, Identif. Guide Vietnam. Orch.:

17; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003,

Slipper Orch. Vietnam: 145. – Cypripedium concolor Bateman, 1865, Curtis’s Bot.

Mag. 91: t. 5513.

Described from Myanmar (“Moulmein”). Type (“cult. Rucker, Parish s.n.”) –

K (holotype), W (isotype).

Lithophytic herb with 4–6 oblong leaves 8–16 by 2–4 cm, distinctly tessellated

above, spotted with purple-violet below. Peduncle erect, 1–2(3) flowered, 5–8 cm tall,

pubescent, dirty-purple spotted. Flowers apple fragrant, 5–7 cm across, white to yellow,

finely sparsely spotted with purple. Pedicel and ovary 3–5 cm long, white pubescent.

Sepal, synsepal and petals sub-similar, ovate to oblong-ovate, 2–4.5 by 2–3 cm. Lip

ellipsoidal to narrowly ovate, fleshy, 2.5–3.8 by 1–1.8 cm. Staminode white with yellow

center, finely purple marked, ovate, toothed to acute at apex, 1–1.3 by 1–1.2 cm wide.

Fig. 18, a-d; 23, i.

Ecology. Broad-leaved, evergreen forests and scrub on rocky limestone. 10–

600 m. Fl. March – June. Not rare (VU).

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Fig. 17. Paphiopedilum micranthum: a – flowering plant, b-d – staminode. P. hangianum: e –

flowering plant, f – staminode. P. emersonii: g – flowering plant, h, i – staminode.


Turczaninowia 2008, 11(1) : 5–168

Distribution. Vietnam (Bac Kan, Bac Giang, Ha Nam, Ha Noi, Hai Phong,

Hoa Binh, Lang Son, Nam Dinh, Ninh Binh, Quang Binh, Quang Ninh, Quang Tri, Son

La, Thai Nguyen, Thanh Hoa, Vinh Phuc), SE Myanmar, S China, Thailand, Laos,

Cambodia?

Studied specimens. Bac Kan, Ba Be, HAL 118 (HN, LE), HLF 615 (HN,

LE); Bac Kan, Cho Don, HLF 841 (HN, LE); Hai Phong, Cat Ba, LX-VN 3393/7

(HN, LE); LX-VN 3747 (HN, LE); Hoa Binh, Mai Chau, DKH 8178 (HN, LE, MO);

Lang Son, Huu Lung, NTH 3259 (HN, K, LE); Ninh Binh, Cuc Phuong, NTH 5383

(HN), NMC 1600 (HN); Quang Binh, Minh Hoa, HAL 5840 (HN), HAL 6037 (HN);

Quang Ninh, Halong Bay, NTH 2631 (HN, LE); Quang Tri, Huong Hoa, HLF 5935

(HN); Son La, Moc Chau, NTH 2922 (HN, LE); Thanh Hoa, Quan Hoa, HAL 3705

(HN), HAL 3757 (HN).

Notes. Common lowland limestone species extinct in many places of its former

occurrence. Color of flowers varies from yellowish-white to bright yellow. Occasional

flowering sometimes observed in December.

8. P. hirsutissimum (Lindl. ex Hook.) Stein, 1892, Orchideenbuch: 470;

Seidenf. 1992, Orch. Indochina: 19; Aver. 1994, Identif. Guide Vietnam. Orch.: 19;

Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003,

Slipper Orch. Vietnam: 156. – Cypripedium hirsutissimum Lindl. ex Hook. 1857,

Curtis’s Bot. Mag. 83: t. 4990.

Described from NE India (“East Indian Plants”). Type (“cult. Parker”) – K.

Distribution. NE India, Myanmar, SW China, N Thailand, N Laos, N Vietnam.

Key to varieties

1. Flowers (10)12–16 cm wide; petals (5)7–8 cm long, in basal half strongly undulate on

margins ............................................................................ 8a. P. hirsutissimum var. esquirolei

– Flowers 6–8(10) cm wide; petals (2.6)3–4(5) cm long, straight or hardly undulate on

margins ........................................................................ 8b. P. hirsutissimum var. chiwuanum

8a. P. hirsutissimum var. esquirolei (Schleichter) Karasawa et Saito, 1982,

Bull. Hiroshima Bot. Gard. 5: 40; Aver. et al. 2003, Slipper Orch. Vietnam: 159. –

P. esquirolei Schleichter, 1919, Repert. Spec. Nov. Regni Veg. Beih. 4: 39. – P. saccopetalum

Hua, 1998, Die Orchidee, 49, 1: 38, forma aberr.

Described from China (“China, Guizhou (Kweichow)”). Type (“Esquirol

3277”) – B (destroyed).

Clustering lithophytic herb, stems with 5–7 linear-ligulate leaves 20–45 by

1.5–2 cm, pure green, few spotted with purple on lower surface near the base. Peduncle

1-flowered, 17–25 cm tall, purple-black pubescent, dirty-purple spotted. Flowers 10–

16 cm wide, glossy. Pedicel and ovary 5–7.5 cm long, densely hairy. Sepal and synsepal

similar, olive-green, heavily suffused with brown, broadly ovate, 3–4.5 by 2–4 cm.

Petals olive-green, rose-purple in apical part, spathulate, 5–8 by 1.5–3 cm, half twisted,

with strongly undulated margins. Lip olive-green, spotted with rose-purple, goblet-

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shaped, 3–5 by 1.5–2.8 cm. Staminode pale purple with glossy olive-brown center,

sub-quadrate, 1 by 0.8 cm. Fig. 18, e-h; 24, a.

Ecology. Broad-leaved evergreen, mixed and coniferous forests on rocky

crystalline limestone. 350–1250 m. Fl. March – May. Not rare (VU).

Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Hoa Binh, Lao Cai,

Ninh Binh, Son La, Thai Nguyen, Thanh Hoa, Tuyen Quang). SW China, N Thailand,

N Laos.

Studied specimens. Bac Kan, Cho Don, HAL 4669 (HN), HLF 770 (HN, LE), HLF

835 (HN, LE); Bac Kan, Na Ri, HAL 4893 (HN), Cao Bang, Ha Lang, CBL 694 (HN, LE), CBL

1677 (HN, LE); Cao Bang, Nguyen Binh, CBL 1628a (LE – photo); Cao Bang, Tra Linh,

Averyanov et al. s.n., 1996 (HN, LE), Averyanov et al. s.n., 1997 (HN, LE), CBL 982 (HN, LE), DH

4349 (HN, LE); Cao Bang, Trung Khanh, HAL 5545 (HN); Ha Giang, Cuan Ba, DKH 6213 (HN,

LE); Ha Giang, Dong Van, NTH 3532 (HN, LE); Hoa Binh, Da Bac, HAL 290 (HN, LE), HAL 318

(HN, LE); Hoa Binh, Mai Chau, DKH 7748 (HN, LE, MO), DKH 8187 (HN, LE, MO); Hoa Binh,

Tung Khe, Averyanov et al. s.n., 1995 (HN, LE); Lao Cai, Sapa, Tachtajan 56 (LE); Ninh Binh,

Cuc Phuong, HAL 1637 (HN), HAL 2854 (HN), DDS 11325 (HN), NMC 933 (HN), NMC 1593

(HN); Son La, Moc Chau, NTH 2971 (HN, LE) HAL 9362 (HN, LE), HAL 9506 (HN, LE), HAL

9410 (HN, LE); Son La, Yen Chau, HAL 9433 (HN, LE); Thai Nguyen, Dong Hy, HAL 28 (HN,

LE); Thanh Hoa, Ba Thuoc, HAL 927 (HN, LE), Thanh Hoa, Ba Thuoc, HAL 969 (HN, LE),

Thanh Hoa, Ba Thuoc, HAL 1077 (HN, LE), Thanh Hoa, Ba Thuoc, HAL 2986 (HN), Thanh

Hoa, Ba Thuoc, HAL 3288 (HN), Thanh Hoa, Ba Thuoc, HAL 4345 (HN); Thanh Hoa, Quan

Hoa, HAL 3716 (HN), HAL 3933 (HN); Tuyen Quang, Na Hang, HLF 663 (HN, LE).

Notes. In the past one of the most common lithophytic species on rocky

limestone formations all over northern Vietnam. Very rarely are observed albino forms

and forms with inflorescence bearing 2 flowers.

8b. P. hirsutissimum var. chiwuanum (T. Tang et F.T. Wang) P.J. Cribb,

1987, Gen. Paphiopedilum: 140; Aver. et al. 2003, Slipper Orch. Vietnam: 166. – P. chiwuanum

T. Tang et F.T. Wang, 1951, Acta Phytotax. Sin. 1: 56.

Described from S China (“China, south-east Yunnan, Foo-ning Hsien, Banloun,

700 m.”). Type (“15 April 1940, Wang 88252”) – PE.

Differs in small flowers, 6–8(10) cm wide. Dorsal sepal 1.8–2.6 by 1.8–2.5

cm. Synsepal 1.8–2.4 by 1.1–1.5 cm. Petals, straight or hardly undulate on margins,

2.6–4(5) by 1–1.3 cm. Lip 1.8–2.6 by 1.2–1.5 cm. Fig. 18, i-k; 24, b.

Ecology. Broad-leaved evergreen, mixed and coniferous forests on rocky

crystalline limestone. 750–1450 m. Fl. March – May. Not rare (VU).

Distribution. Vietnam (Cao Bang, Hoa Binh, Son La), S China.

Studied specimens. Cao Bang, Bao Lac, CBL 1370 (HN, LE), CBL 1588 (HN, LE);

Hoa Binh, Da Bac, HAL 319 (HN, LE); Hoa Binh, Mai Chau, HAL 624 (HN, LE); Hoa Binh, Tung

Khe, Averyanov et al. s.n., 1995 (LE); Son La, Yen Chau, DKH 7260 (HN, LE).

Notes. Montane race closely allied to previous variety.

9. P. barbigerum T. Tang et F.T. Wang, 1940, Bull. Fan. Mem. Inst. Bot. Ser.

10: 23; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al.


Turczaninowia 2008, 11(1) : 5–168

Fig. 18. Paphiopedilum concolor: a – flowering plant, b-d – staminode; P. hirsutissimum var.

esquirolei: e – flowering plant and flower, f-h – staminode. P. hirsutissimum var. chiwuanum: i –

flower, j, k – staminode. P. barbigerum var. coccineum: l – flowering plant, m – staminode.

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2003, Slipper Orch. Vietnam: 170. – P. insigne (Wall. ex Lindl.) Pfitz. var. barbigerum

(T. Tang et F.T. Wang) Braem, 1988, Paphiopedilum: 113. – P. × areeanum Gruss,

2001, Die Orchidee, 52, 5: 117. – P. rhizomatozum S.C. Chen et Z.J. Liu, 2002, Journ.

Wuhan Bot. Res. 20, 1: 12.

Described from S China (“China, Kweichow (Guizhou)”). Type (“Cavalerie

& Fortunat 1794”) – PE (holotype), P (isotype).

Distribution. N Myanmar, N Thailand, S China, N Vietnam.

Key to varieties

1. Median sepal white with uniformly pale pink, pink, pink-brownish or carmine center ...

................................................................................................. 9a. P. barbigerum var. cocineum

– Median sepal white or yellowish-pink with pink to pink-carmine center spotted with

numerous contrast dark brown to brown-purple blotches ... 9b. P. barbigerum var. aspersum

9a. P. barbigerum var. coccineum (Perner et R. Herrmann) W. Cavesrto,

2001, Gen. Paphiopedilum: 200. – P. coccineum Perner et R. Herrmann, 2000, Die

Orchidee, 51, 5: 623. – P. barbigerum var. lockianum Aver. 2002, Komarovia, 2: 11;

Aver. et al. 2003, Slipper Orch. Vietnam: 170. – P. vejvarutianum Gruss et Roellke,

2003, Die Orchidee, 54, 1: 59.

Described from N Vietnam (“Provinz Cao Bang” – obviously wrong).

Lectotype – Ic. “Paph. coccineum Perner & Hermann”, 2000, Die Orchidee, 51, 5:

622, 623, 624. Herbarium authentic materials (“Giay Giot Thieu ohne Nummer”) newer

been located.

Lithophytic herb with 4–6 narrowly lanceolate, uniformly green leaves 8–14

by 0.7–1.2 cm. Peduncle 1-flowered, erect, 12–18 cm long, shortly brownish pubescent.

Flower 6–10 cm across. Pedicel and ovary 3.1–4 cm long, shortly light brown pubescent.

Median sepal white with pink, carmine or carmine-brownish center, broadly ovate, 3–

5 by 2.6–4.2 cm. Synsepal elliptic, 2.4–5 by 1.3–2.2 cm. Petals yellowish-brown to

carmine-brown with yellowish margin, spathulate, 3.6–5 by 1–2.2 cm, with undulate

margin. Lip yellowish-brown, yellow-pinkish to carmine-brown, 3–4.5 by 3–3.5 cm.

Staminode light yellow with orange umbo, obovate, 9–11 by 8–10 mm. Fig. 18, l, m;

24, c.

Ecology. Broad-leaved evergreen forests on rocky crystalline limestone. 1000–

1100 m. Fl. September – November. Very rare (CR).

Distribution. Vietnam (Son La), N Thailand?

Studied samples. Son La, Moc Chau, P-9894 (HN), P-10546 (HN, LE), HAL 9406

(HN, LE).

Notes. Variable species, particularly in size and coloration of flower. It is one of the

rarest slipper orchid species in northern Vietnam. Occurrence in Thailand (P.vejvarutianum)

was reported on the base of cultivated plants origin of which needs confirmation.

9b. P. barbigerum var. aspersum (Aver.) Aver. comb. et stat. nov. –

P. × aspersum Aver. 2002, Komarovia, 2: 17; Aver. et al. 2003, Slipper Orch. Vietnam:


Turczaninowia 2008, 11(1) : 5–168

254. – P. trantuanhii Gaetan Gogniat et X.G. de Loubresse, 2008, SOS Bull. 1, 1: 2.

Described from N Vietnam (“Son La prov., Moc Chau distr., …”). Type

(“Phan Ke Loc, P-9895”) – HN.

Differs from previous variety in median sepal with pink to pink-carmine center

spotted with numerous contrast dark brown to brown-purple blotches. Fig. 24, d.

Ecology. Broad-leaved evergreen forests on rocky crystalline limestone. 1000–

1100 m. Fl. September – November. Very rare (CR).

Distribution. Vietnam (Son La). Endemic.

Studied samples. Son La, Moc Chau, P-9895 (HN), P-10547 (HN).

Notes. Like previous variety, this is one of the rarest slipper orchid species

with very restricted range. It has intermediate morphology between P. insigne and

other Indochinese races of P. insigne complex (like P. barbigerum var. coccineum

and P. henryanum). Both varieties may be certainly found in limestone regions of

northern Laos allied to Vietnamese border.

10. P. tranlienianum Gruss et Perner, 1998, Caesiana, 11: 66; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam:

175. – P. caobangense Nguyen Thien Tich, 1999, Hoa Canh (1999), 1: 14, nom. illeg.

Described from N Vietnam (“Bac Thai Province, October 1998…” – highly

doubtful). Type (“fl. in cult. November 1998, leg. O. Gruss”) – RO.

Lithophytic herb with 3–6 oblong lanceolate, glossy green leaves, to 18 by

1.7 cm, with paler margin. Peduncle suberect, 1-flowered, 10–18 cm tall, bright green,

yellow to purple-brown hairy. Flower 5.5–6 cm across. Pedicel and ovary 2.9–5 cm

long, green, brown to purple-brown hairy. Dorsal sepal white, with purple veins, suborbiculate,

2.5–3.2 by 2.8–3.4 cm. Synsepal pale green, ovate, 2.4–2.6 by 1.1–1.8 cm.

Petals olive-green, veined with chocolate-brown, oblong, 3–3.4 by 0.7–0.9 cm, glossy,

white-ciliate, strongly undulate along margins. Lip olive-green, glossy, with a prominent

claw-like base, 3.7–3.9 by 1.6 cm. Staminode yellow, obcordate to obovate, 8–10 by

7–9 mm, with glossy umbo. Fig. 19, a-e; 24, e.

Ecology. Broad-leaved, evergreen forests on rocky crystalline and slaty

stratified. 400–750 m. Fl. September – November. Very rare (CR).

Distribution. N Vietnam (Thai Nguyen, Tuyen Quang). Endemic.

Studied samples. Thai Nguyen, Dong Hy, HAL 30 (HN); Tuyen Quang, Na Hang,

HAL 183 (HN).

Notes. Local endemic with very restricted distribution. The plants exhibit

essential variation in size of flower. Very rare they develop inflorescence with 2 flowers.

11. P. helenae Aver. 1996, Bot. Journ. (Leningrad) 81, 9: 108; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam:

182. – P. helenae f. aureum Gruss et Rоth, 1999, Die Orchidee, 50, 1: 3. – P. delicatum

Z.J. Liu et J.Y. Zhang, 2001, Acta Phytotax. Sin. 39, 1: 78.

Described from N. Vietnam (“Cao Bang Prov., Tra Linh Distr., Quoc Toan

Subdistr., 850–900 m, near Thang Hen Lake, environs of Thang Hen and Lung Tao

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villages”). Type (“21 October 1995, L. Averyanov, N.T. Hiep, D.D. Huyen CB 12”) –

LE (holotype), HN (isotype).

Dwarf lithophytic herb with 3–5 coriaceous to rigid, rather succulent, oblong

leaves 3–12 by 0.6–2 cm, green with yellowish margin. Peduncle erect to almost

horizontal, one-flowered, 4–7 cm long, shortly black-purple pubescent. Flower 2.5–6

cm across. Pedicel and ovary 2–3.5 cm long, shortly black-purple pubescent. Median

sepal dull to bright yellow with white margin, obovate to circular, 1.8–3.5 by 1.5–3 cm.

Synsepal white, ovate, 1.5–2.5 by 0.8–1.5 cm. Petals pale yellow to bright orange

with darker nerves, straight, ligulate, 1.5–3.5 by 0.4–0.8 cm. Lip pale pink to brownishorange,

saccate, 2–3 by 1.5–2 cm. Staminode yellow, obovate to circular, 7–8 mm

across, with central bright yellow to greenish umbo. Fig. 19, f-h; 24, f.

Ecology. Broad-leaved evergreen, mixed and coniferous dry forests on rocky

crystalline limestone. 500–900 m. Fl. September – November. Rare (EN).

Distribution. Vietnam (Bac Kan, Cao Bang, Lang Son), S China?

Studied samples. Bac Kan, Na Ri, HAL 4981 (HN); NTH 3716 (HN, LE), DKH

7539 (HN, LE, MO); Cao Bang, Ha Lang, CBL 648 (HN, LE), CBL 744 (HN, LE), CBL 767/1

(LE), CBL 789 (HN, LE), CBL 1690b (LE); Cao Bang, Thach An, CBL 830 (HN, LE), CBL 911

(HN, LE); Cao Bang, Tra Linh, CB 12 (HN, LE), CBL 1140 (HN, LE), CBL 1222/1 (LE); Cao

Bang, Trung Khanh, HAL 5456 (HN), HAL 5628 (HN); Lang Son, Bac Son, HAL 6728 (HN).

Notes. Local endemic with very restricted distribution. Quite possible this species

occurred in the past in border territory of S China allied to Cao Bang and Lang Son

provinces of Vietnam. It is very variable in plant dimensions and flower coloration. Flowers

vary from very small, about 2.5 cm wide, to relatively large up to 6–7 cm across.

12. P. henryanum Braem, 1987, Schlechteriana, 1, 1: 4; Seidenf. 1992, Orch.

Indochina: 19; Aver. 1994, Identif. Guide Vietnam. Orch.: 20; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam:

191. – P. dollii Lоckel, 1987, Die Orchidee, 38, 5: 266. – P. chaoi Hua, 1999, Die

Orchidee, 50, 5: 495.

Described from China (“China, border region to Vietnam”). Type (“1987,

Azadehdel s.n.”) – SCHLE.

Lithophytic herb with 3–6 lanceolate leaves 10–17 by 1.2–1.6 cm, glossy dark

green, with yellowish line along margin. Peduncle suberect to arcuate-pendent, 1flowered,

12–15 cm long, green, purple-brown pubescent. Pedicel and ovary 3.6–

4cm long, light green. Flower 4–6 cm across. Median sepal and synsepal light yellowishgreen

with maroon spots, broadly-ovate to subcircular; sepals 3.3–3.6 by 2.9–3.5 cm,

undulate on margins; synsepal 2.5–2.8 by 1.6 cm. Petals dull yellowish to pink-violet,

spotted with maroon, spreading, oblong-spathulate, 3.4–3.6 by 1.4–1.6 cm, with undulate

margins. Lip rose to pink-violet, deeply saccate, 3.7–4.3 by 1.9–2.1 cm. Staminode

yellow with purple tint, broadly obovate, 7–10 by 8–9 mm. Fig. 19, i-l; 24, g.

Ecology. Broad-leaved, mixed and coniferous, evergreen or semi-deciduous forests

on rocky crystalline limestone. 700–1400 m. Fl. September – November. Rare (EN).

Distribution. Vietnam (Bac Kan, Ha Giang), S China.


Turczaninowia 2008, 11(1) : 5–168

Fig. 19. Paphiopedilum tranlienianum: a – flowering plant, b – flower, c-e – staminode; P. helenae:

f – flowering plant, g – flower, h – staminode; P. henryanum: i – flowering plant, j – flower, k, l–

staminode.

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Studied samples. Bac Kan, Cho Don, HAL 4767 (HN), HAL 4817 (HN), HLF 772

(HN, LE), HLF 805 (HN, LE); Ha Giang, Dong Van, NTH 3533 (HN, LE); Ha Giang, Meo Vac,

CBL 1875 (HN, LE), NTH 3323 (HN, LE), HAL 8497 (HN); Ha Giang, Quan Ba, NTH 3590 (HN,

LE), DKH 6159 (HN, LE); Ha Giang, Vi Xuyen, DKH 6460 (HN, LE).

Notes. Local endemic with very restricted distribution. Very variable species,

particularly in size of flowers, form and spotting of tepals. Rarely are observed albino

specimens.

13. P. gratrixianum Masters ex Rolfe, 1905, Orchid Rev. 13: 63; Seidenf.

1992, Orch. Indochina: 17; Aver. 1994, Identif. Guide Vietnam. Orch.: 20; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003, Slipper Orch.

Vietnam: 199. – Cypripedium gratrixianum Masters, 1905, Gard. Chron. Ser. 3, 37:

76, fig. 35, nom. inval., non B.C. Williams, 1897. – P. villosum (Lindl.) Stein var.

gratrixianum (Masters ex Rolfe) Braem, 1988, Paphiopedilum: 119.

Described from Laos (“Native of Annam”). Type (“Micholitz s.n., cult. Sander”)

– K.

Distribution. S Laos, Vietnam.

Key to varieties

1. Median sepal with green, olive-green to yellowish-green center, sometimes slightly

flushed with purple-violet, spotted with numerous small spots ...........................................

.................................................................................... 13a. P. gratrixianum var. gratrixianum

– Median sepal pure white, occasionally with light green or purplw violet tint at the

base, spotted with few relatively large blotches .......... 13b. P. gratrixianum var. daoense

13a. P. gratrixianum var. gratrixianum.

Lithophytic or terrestrial herb with 4–7 lanceolate, pure green, leaves up to 30

by 2–3 cm, spotted with dirty purple-violet on underside near the base. Peduncle 1flowered,

erect, up to 25 cm long, green, brownish-purple hairy. Pedicel and ovary 4–

5.5 cm long, pale green. Flower 7–8 cm across. Median sepal with green to brownishgreen

center and white margin, more or less heavily spotted with brown-purple, broadly

ovate to subcircular, 4.8–5.2 by 4.4–4.6 cm. Synsepal pale green, ovate-elliptic, 3.5–

5 by 2.5 cm. Petals yellow-brown, veined with purple-brown, glossy, spreading, spathulate,

4.5–5.2 by 1.8–2.5 cm, margins undulate or reflexed. Lip yellow to yellow-brown,

glossy, 4–4.2 by 2.4–2.8 cm. Staminode yellow, obovate, 10–12 mm long and wide,

shortly acute at the apex. Fig. 24, h.

Ecology. Broad-leaved, evergreen forests on granite, gneiss and similar silicate

rocks. 900–1500 (2000?) m. Fl. October – December. Very rare (DD).

Distribution. Vietnam (Lam Dong), S Laos.

Studied samples. Lam Dong, cult., Averyanov s.n., 2004 (LE – photo), Averyanov

s.n., 2005 (LE – photo).

Notes. Available reports of this variety for Vietnam are based on cultivated

plants supposedly collected in montane regions along Laos-Vietnamese border.


Turczaninowia 2008, 11(1) : 5–168

Occurrence in Vietnam needs confirmation. Very variable variety in coloration of

median. Easily forms hybrids with P. villosum.

13b. P. gratrixianum var. daoense Aver. 2002, Orch. Rev. 110, 1247: 287.

Described from N Vietnam (“Vinh Phuc Prov., Tam Dao Distr., vicinities of

Tam Dao town (21 о 27’N 105 о 39’E). Tam Dao ridge at an altitude of 950–1000 m. …”).

Type (“10 November 1998. N.T. Hiep, Averyanov L. s.n.”) – HN (holotype.), LE

(isotype).

Differs from type variety in pure white median sepal, rarely light green or

purple violet tinted at the base, spotted with few relatively large purple-brown blotches.

Leaves and inflorescence shorter and all plant more compact and miniature. Fig. 20, ac;

24, i.

Ecology. Broad-leaved, evergreen humid forests on granite, rhyolite and similar

silicate rocks. 900–1500 m. Fl. October – December. Very rare (EN).

Distribution. Vietnam (Lai Chau, Lao Cai, Thai Nguyen, Vinh Phuc). Endemic.

Studied samples. Lao Cai, Sa Pa, Averyanov s.n. 2005 (LE – photo); Vinh Phuc,

Tam Dao, 10 November 1998. N.T. Hiep, Averyanov L. s.n. (HN, LE), VH 5041 (HN, LE).

Notes. According to observations not confirmed by herbarium collections,

this variety may be rarely found also on rocky limestone.

14. P. villosum (Lindl.) Stein, 1892, Orchideenbuch: 490; Seidenf. 1992, Orch.

Indochina: 19; Aver. 1994, Identif. Guide Vietnam. Orch.: 19; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch. Vietnam:

208. – Cypripedium villosum Lindl. 1854, Gard. Chron. (1854): 135.

Described from Myanmar (“Burma”). Type (“Lobb s.n.”) – K.

Distribution. NE India, Myanmar, S China, Laos, Vietnam, Thailand.

Key to varieties

1. Median sepal broadly oblanceolate to narrowly obovate, boldly spotted with deep

brown fused blotches, with strongly reflexed lateral margins throughout its length;

petals often marked with purple-brown, narrowly spatulate, distinctly attenuate to

narrowing stipitate base ............................................................ 14e. P. villosum var. boxallii

– Median sepal obovate to suborbicular, without distinct spots, with strongly reflexed

lateral margins at lower part; petals without purple-brown marks, with more or less

broad, not distinctly stipitate base ................................................................................... 2

2. Median sepal dark brown with more or less wide yellowish-green to whitish margin ....

.................................................................................................... 14a. P. villosum var. villosum

– Median sepal light yellowish-green, dull pink to purple, or white with more or less

wide median longitudinal brown stripe ............................................................................ 3

3. Median sepal white, with more or less wide median longitudinal brown stripe,

sometimes with broad dark brown central area occasionally margining with green .......

................................................................................................ 14b. P. villosum var. annamense

– Central area of median sepal dull pink to dark purple or dull yellowish-green tinted

with light brown along midvein ........................................................................................ 4

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

4. Central area of median sepal dull pink to dark purple, occasionally greenish at the

base; sepal margin commonly white ............................ 14d. P. villosum var. fusco-roseum

– Median sepal dull yellowish-green, tinted with light brown along midvein, occasionally

with white or yellowish-white margin .......................... 14c. P. villosum var. fusco-viride

14a. P. villosum var. villosum.

Epiphytic herb with 4–5, ligulate, pure green leaves, 14–42 by 2.5–4 cm, on

beneath, purple-spotted at the base. Peduncle suberect to pendent, 1-flowered, 7–

24 cm long, green, whitish to black-purple villose. Pedicel and ovary, 3–6 cm long,

light-green, densely villose. Flower 7.5–13.5 cm across. Dorsal sepal dark brown with

yellowish-green to whitish margin, obovate, 4.5–7 by 3–4.6 cm. Synsepal pale green,

narrowly ovate, 3.8–7.6 by 1.8–2.6 cm. Petals yellow- to reddish-brown, often with a

central maroon stripe, glossy, spathulate, 4.7–8.6 by 2.5–4.6 cm. Lip yellow-brown

flushed with reddish, 4–6.8 by 3–3.8 cm. Staminode yellow, obovate, about 16 by 14

mm, with a central glossy umbo. Fig. 25, a.

Ecology. Shady broad-leaved evergreen and mixed humid forests mainly on

silicate rocks. 1300–2000 m. Fl. November – March. Rare (VU).

Distribution. Vietnam (Dak Nong, Lam Dong), NE India, Myanmar, SW

China, Laos, Thailand.

Studied samples. Dak Nong, Dak Glong, HLF 5671 (HN); Lam Dong, Dalat,

Averyanov s.n., 2005 (LE – photo).

Notes. This is most common and widespread variety. It commonly grows as

typical humus epiphyte.

14b. P. villosum var. annamense Rolfe, 1907, Curtis Bot. Mag. 133: t. 8126;

Aver. et al. 2003, Slipper Orch. Vietnam: 215.

Described from S Vietnam (“Annam, Micholitz s.n., cult. Sander”). Lectotype –

Ic. “P. villosum var. annamense” (1907, Curtis’s Bot. Mag. 133, tab. 8126).

This variety differs from the type in pure white median sepal with narrow

chestnut-brown longitudinal stripe. Occasionally this central brown stripe on sepal

becomes wider forming broad chestnut to brown-purple central area margining with

green. Fig. 20, d-f; 25, b.

Ecology. Shady broad-leaved mixed and evergreen humid forests mainly on

silicate rocks. 1300–2000 m. Fl. November – March. Occasional (VU).

Distribution. Vietnam (Dak Nong, Khanh Hoa, Lam Dong). Endemic.

Studied samples. Dak Nong, Dak Glong, HLF 5671 (HN); Gia Lai, Chu Pah, Phan

Ke Loc, P-9863 (HN); Khanh Hoa, Khanh Son, VH 4349 (HN, LE); Lam Dong, Dalat, Averyanov

s.n., 1995 (LE – photo), Averyanov s.n., 2005 (LE – photo), O. Zaitsev et al. 153 & 281 (LE); Lam

Dong, Lac Duong, VH 3120 (HN, LE), VH 3333 (HN, LE).

Notes. Eastern vicarious race of the species, which is rather common in

montane forests of southern Vietnam. Very variable. Grows commonly as humus

epiphyte.


Turczaninowia 2008, 11(1) : 5–168

14c. P. villosum var. fusco-viride Aver. 2002, Komarovia, 2: 17.

Described from S Vietnam (“Gia Lai prov., Chu Pah distr.”). Type (“27.05.2001,

Phan Ke Loc, P-9845B”) – HN.

Flowers wholly dull yellowish with dull yellowish-green median sepal hardly

tinted with brown at the center. It commonly grows as lithophytic or terrestrial herb.

Fig. 25, c.

Ecology. Shady broad-leaved mixed and evergreen humid forests mainly on

silicate rocks. 1300–1700 m. Fl. October – December. Very rare (EN).

Distribution. Vietnam (Lam Dong, Gia Lai). Endemic.

Studied samples. Gia Lai., Chu Pah, P-9845B (HN); Lam Dong, Dalat, Averyanov

s.n., 2005 (LE – photo).

Notes. Local endemic of eastern Indochina. May be found on territories of

Cambodia and Laos allied to Vietnamese border.

14d. P. villosum var. fusco-roseum Aver. 2002, Komarovia, 2: 17. – P. macranrhum

Z.J. Liu et S.C. Chen, 2002, Acta Bot. Yunnan. 24, 6: 712.

Described from S Vietnam (“Gia Lai prov., Chu Pah distr.”). Type (“27.05.2001,

Phan Ke Loc, P-9845A”) – HN.

Median sepal with pink to purple center bordering with pure. Occasionally

pink central area of the sepal at the base tinted with light green. Often whole flower

may be light dull yellowish with nearly white median sepal slightly tinted with pink. Fig

25, d.

Ecology. Shady broad-leaved mixed and evergreen humid forests mainly on

silicate rocks. 1300–1700 m. Fl. October – December. Very rare (EN).

Distribution. Vietnam (Lam Dong, Gia Lai), E Cambodia.

Studied samples. Gia Lai, Chu Pah, P-9845A (HN); Lam Dong, Dalat, Averyanov

s.n., 2005 (LE – photo).

Notes. Local endemic of eastern Indochina. May be also found on the territory

of Laos allied to Vietnamese border. It commonly grows as lithophytic and terrestrial.

14e. P. villosum var. boxallii (Reichenb. f.) Pfitz. 1903, in Engler, Pflanzenr.

4, 50, 12, Orch. Pleon.: 73; Aver. et al. 2003, Slipper Orch. Vietnam: 217. –

Cypripedium boxallii Reichenb. f. 1877, Gard. Chron. n. s. 7: 367.

Described from Myanmar (“Burma, Moulmein”). Type (“Boxall s.n.”) – W.

This variety is distinguished by its deep colored flowers with a boldly spotted narrow

median sepal and narrow lateral petals heavily striped with purple-brown. Fig. 25, e.

Ecology. Shady broad-leaved evergreen humid mountain forests? Fl.? Probably

very rare (DD).

Distribution. Vietnam (Lao Cai, Lai Chau)?, Myanmar.

Studied samples. Lao Cai, Sa Pa, Averyanov s.n., 2004 (LE – photo).

Notes. Many plants of this rare variety collected in nature are cultivated in Sa

Pa town. Meanwhile the occurrence of this plant in Vietnam needs confirmation by

herbarium collections.

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15. P. appletonianum (Gower) Rolfe, 1896, Orchid Rev. 4: 364; Seidenf.

1992, Orch. Indochina: 19; Aver. 1994, Identif. Guide Vietnam. Orch.: 21; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003, Slipper Orch.

Vietnam: 218. – Cypripedium appletonianum Gower, 1893, The Garden, 1893: 95. –

P. hainanense Fowlie, 1987, Orchid Dig. 51, 2: 69. – P. cerveranum Braem, 1999,

Orchid. Cult. Protect. 38: 28. – P. tridentatum S.C. Chen, Z.J. Liu et J.Y. Zhang,

2001, Acta Phytotax. Sin. 39, 5: 455.

Area of the type origin is unknown (“a plant was introduced with some plants

of C. Hookerae”). Type (“hort. F. Appleton, April 1893”) – K.

Terrestrial or lithophytic herb with 4–8 oblong-elliptic leaves 7–25 by 2–4 cm,

distinctly tessellated. Peduncle erect, 1(2)-flowered, 20–50 cm long, purple, shortly

pubescent. Pedicel and ovary dull dirty purple spotted, 3–6 cm long. Flower 6–12 cm

across. Dorsal sepal olive-green, often brown-purple striped, ovate, 2.7–4.5 by 2–

3.2 cm. Synsepal green, elliptic, 2–3 by 1.1–1.5 cm. Petals green with darker green

and olive-brown stripes and maroon-black spots in basal half, pale pink-purple to the

apex, spatulate, 4.4–6 by 1.2–1.8 cm, often with marginal maroon-black warts along

upper margin. Lip olive-green, often with pale purple tint, 3.6–4.6 cm long; front margin

more or less distinctly 2–6-dentate. Staminode yellowish to green with yellowish margin

and central yellowish spot, from broadly obcordate to deeply transversely lunate, with

2, 4 or 5 toothed apex, 7–9 by 8–9 mm. Fig. 20, g-k; 25, f.

Ecology. Shady broad-leaved evergreen, mixed and coniferous humid forests

on granite, gneiss and sandstone. 700–2000 m. Fl. March – May. Occasional (VU).

Distribution. Vietnam (Dak Lac, Dak Nong, Khanh Hoa, Kon Tum, Lam

Dong, Ninh Thuan, Quang Binh, Quang Tri, Thua Thien – Hue), China (Hainan),

Thailand, Laos, Cambodia.

Studied samples. Dak Lac, Krong Bong, VH 6351 (HN, LE); Dak Nong, Dak Glong,

HLF 5671 (HN); Khanh Hoa, Khanh Le, QD-12 (LE); Khanh Hoa, Khanh Son, VH 4281 (HN,

LE); Kon Tum, VH 964 (HN, LE); Kon Tum, Kon Plong, VH 5316 (HN, LE); Lam Dong, Dalat,

L. Averyanov, N.T. Hiep VH s.n., 1995 (LE); Lam Dong, Lac Duong, VH 3405 (HN, LE,), VH

4123 (HN, LE); Ninh Thuan, Ninh Son, VH 3530 (HN, LE), VH 3626 (HN, LE); Quang Tri,

Huong Hoa, HLF 5800 d-EXSICCATES OF VIETNAMESE FLORA 0072/HLF5800 (HN);

Thua Thien – Hue, A Luoi, HAL 7607 (HN); Thua Thien - Hue, Phu Loc, HLF 959 (HN), HLF

1049 (HN); Quang Binh, Minh Hoa, HAL 11715 (HN).

Notes. One of the most common and variable species, particularly in coloration

of flowers, form of tepals and staminode. Very rarely some specimens in wild populations

form two-flowered inflorescence.

16. P. callosum (Reichenb. f.) Stein, 1892, Orchideenbuch: 457; Seidenf.

1992, Orch. Indochina: 20; Aver. 1994, Identif. Guide Vietnam. Orch.: 21; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al. 2003, Slipper Orch.

Vietnam: 230. – Cypripedium callosum Reichenb. f. 1886, Gard. Chron. Ser. 2, 26:

326. – P. callosum var. angustipetalum Guillaum. 1924, Bull. Soc. Bot. France Ser.


Turczaninowia 2008, 11(1) : 5–168

Fig. 20.Paphiopedilum gratrixianum var. daoense: a – flowering plant, b, c – staminode;

P. villosum var. annamense: d – flowering plant, e, f – staminode; P. appletonianum: g –

flowering plant, h – flower, i, j – staminode, k – variation of staminode shape.

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

4, 24: 551. – P. amabile auct. non Hallier.: Seidenf. 1975, Contrib. Revis. Orch. Fl.

Cambod. Laos Vietnam: 88; Aver. 1988, Prelim. List Vietnam. Orch. 2: 31.

Described from Thailand (“Siam”). Type (“Regnier s.n.”) – W.

Distribution. Thailand, Vietnam, Laos, Cambodia.

Key to varieties

1. Petals strongly sigmoid, distinctly down deflexed, usually more than 5 cm long,

commonly with ciliate warts along upper and lower margin; median sepal usually

more than 5 cm wide ................................................................. 16a. P. callosum var. callosum

– Petals hardly sigmoid or straight, borne at an angle of 45° to the horizontal, usually

less than 5 cm long, with few ciliate warts along upper margin or without marginal

warts at all; median sepal usually less than 5 cm wide .................................................... 2

2. Petals bear few warts only on the upper margin ......... 16b. P. callosum var. warnerianum

– Petals lack the marginal warts at all ........................ 16c. P. callosum var. potentianum

16a. P. callosum var. callosum.

Terrestrial or lithophytic herb with 3–6 oblong-elliptic, distinctly tessellated

leaves, 10–20 by 3.2–4.8 cm. Peduncle 1-flowered, 20–40 cm tall, marked with purple,

purple-pubescent. Pedicel and ovary 3–6.5 cm long, green, flushed with purple. Flowers

8–11 cm across. Dorsal sepal white, veined with purple and green, with small silver

warts near the base, broadly obovate, 4–5.5 by 4.2–6 cm. Synsepal white to greenish,

elliptic, 2.7–3.2 by 1.6–2.5 cm. Petals greenish with a purple apical third, spotted with

maroon, sigmoid, ligulate, 4.6–6.8 by 1.2–1.8 cm, with maroon ciliated marginal warts.

Lip dull brown-purple, warted on side lobes, 2.5–4.4 by 2–2.5 cm. Staminode white to

yellowish, with dark green veins, lunate, 7 by 11 mm. Fig. 21, a-c; 25, g.

Ecology. Shady evergreen broad-leaved forests mainly on granite and

sandstone. 300–1300 m. Fl. April – June. Very rare (EN).

Distribution. Vietnam (Dak Nong, Gia Lai, Khanh Hoa, Kien Giang, Lam

Dong, Quang Tri), Thailand, Laos, Cambodia.

Studied samples. Dak Nong, Dak Glong, HLF 5575 (HN); Gia Lai, Chu Pah, P.K.

Loc P-9864 (HN); Khanh Hoa, Tourane, Delacour s.n. (P); Kien Giang, Phu Quoc, N.V. Khoi

s.n. (LE – photo); Lam Dong, Dalat area., Grillet 51 (P), Grillet 215 (P), Averyanov, Hiep VH

s.n., 2000 (LE); Quang Tri, Lao-bao, Dalacour s.n. (P).

Notes. Widespread, but presently very rare and highly endangered species.

Obviously old records for Vietnam were based on populations which are now already

extinct. Very variable, particularly in form and coloration of sepal and petals. Some

specimens in the wild rarely form 2-flowered stalk as well as albino or sub-albino flowers.

16b. P. callosum var. warnerianum (T. Moore) P.J. Cribb ex Aver. comb.

nov. – Cypripedium barbatum var. warnerianum T. Moore, 1878, Warner Select.

Orchid. Ser. 3, Pl. 3, t. 11. – C. callosum var. sublaeve Reichenb. f. 1888, Gard.

Chron. Ser. 3, 3: 331. – P. callosum var. sublaeve (Reichenb. f.) P.J. Cribb, 1987,

Gen. Paphiopedilum: 188.

Origin of the type unknown. Lectotype – Ic. “Paphiopedilum callosum var.


Turczaninowia 2008, 11(1) : 5–168

warnerianum” (T. Moore, 1878, l.c. Pl. 3, t. 11).

Differs from type variety in smaller flowers. Median sepal smaller, 3.4–4.2 by

3–4.1 cm. Petals shorter, relatively broader, more or less spread or hardly sigmoid,

with warts only on the upper margin, usually uniform green to pink-purple with more

dark longitudinal nerves, rarely spotted. Fig. 25, h.

Ecology. Shady evergreen broad-leaved lowland and submontane forests. Fl.

April – June. Very rare (DD).

Distribution. Vietnam (Lam Dong), S Thailand, NW Malacca.

Studied samples. Lam Dong, Dalat, Averyanov, HAL s.n. 2004 (LE – photo).

Notes. This taxon having main area of distribution in Peninsular Thailand and

adjacent north-west Malacca is sometimes regarded as natural hybrid between two

vicarious species – P. callosum (distributed in mainland Southeast Asia) and P. barbatum

(from Malacca Peninsular and northern Sumatra). Rarely occur samples with

2-flowered inflorescence, as well as sub-albino and albino forms.

16c. P. callosum var. potentianum (O. Gruss et J. Roeth) P.J. Cribb, 1998,

Gen. Paphiopedilum, ed. 2: 337. – P. potentianum O. Gruss et J. Roeth, 1995, Caesiana,

5: 39.

Described from Thailand (“Thailand”). Type (“cult. E. et G. Potent, Roth

s.n.”) – HAL.

The variety differs from the type in unspotted petals lacking marginal warts

on the upper margin and in its narrower dorsal sepal, about 4 by 2.6 cm. Fig. 25, i.

Ecology. Shady evergreen broad-leaved lowland and submontane forests. Fl.

April – June. Very rare (DD).

Distribution. Vietnam (Lam Dong), Thailand.

Studied samples. Lam Dong, Dalat, Averyanov, HAL s.n. 2004 (LE – photo).

Notes. May be certainly found in lowlands of Laos and Cambodia. Rarely

forms sub-albino and albino forms.

17. P. purpuratum (Lindl.) Stein, 1892, Orchideenbuch: 481; Seidenf. 1992,

Orch. Indochina: 20; Aver. 1994, Identif. Guide Vietnam. Orch.: 21; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 49; Aver. et al. 2003, Slipper Orch.

Vietnam: 236. – Cypripedium purpuratum Lindl. 1837, Bot. Reg. 23: t. 1991. –

C. sinicum Hance ex Reichenb. f. 1853, Walp. Ann. 3: 602. – P. sinicum (Hance)

Stein, 1892, Orchideenbuch: 481. – P. purpuratum (Lindl.) Stein var. hainanense

F.Y. Liu et Perner, 2001, Orchidee, 52, 1: 64. – P. aestivum Z.J. Liu et J.Y. Zhang,

2001, Acta Phytotax. Sin. 39, 6: 568, fig. 1.

Described from Hong Kong? Lectotype – Ic. “Cypripedium purpuratum”

(Lindl. 1837, Bot. Reg. 23, t. 1991).

Terrestrial herb with 4–8 oblong-elliptic, distinctly tessellated leaves, 7–17 by

2.3–4.2 cm. Peduncle 1-flowered, 10–20 cm tall, purple-violet, deep purple pubescent.

Pedicel and ovary 2.5–4 cm long, green, whitish pubescent. Flowers 7–10 cm across.

Dorsal sepal white, greenish at the base, heavily veined with purple broadly cordate,

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

2.5–3.5 by 2.2–4.3 cm. Synsepal green with darker veins, narrowly ovate, 2–3.5 by

1.2–1.6 cm. Petals glossy deep purple-maroon, heavily spotted with black-maroon

spots, spreading, narrowly elliptic, 3.5–4.6 by 0.9–1.3 cm. Lip purple, 3.3–4.4 by 2–

2.7 cm, verrucose on incurved side-lobes. Staminode yellowish, purple tinted along

margin, with green veins, lunate, with narrow falcate acute side lobes, 8 by 8–11 mm.

Fig. 21, d-g; 26, a.

Ecology. Broad-leaved evergreen and semi-deciduous forests on highly eroded

crystalline limestone at elev. 600–1400 m. Fl. September – October. Very rare (EN).

Distribution. Vietnam (Cao Bang, Lao Cai, Tuyen Quang), SE China.

Studied samples. Cao Bang, Bao Lac, CBL 1458b (LE); Lao Cai, Sa Pa, P.K. Loc

s.n., 2000 (HN, LE – photo); Tuyen Quang, Na Hang, HLF 662 (HN, LE).

Notes. Rare species, which populations commonly include very few scattered

samples. Leaf tessellation varies from whitish or pale green to deep green. Rarely

forms 2-flowered inflorescence.

18. P. dianthum T. Tang et F.T. Wang, 1940, Bull. Fan Mem. Inst. Biol. Ser.

10: 24; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 48; Aver. et al.

2003, Slipper Orch. Vietnam: 244. – P. parishii (Reichenb. f.) Stein var. dianthum

(T. Tang et F.T. Wang) Karasawa et Saito, 1982, Bull. Hiroshima Bot. Gard. 5: 38.

Described from S China (“China, Yunnan, Mengtze, …”). Type (“Wang et

Liu, 83446”) – PE.

Large lithophytic herb forming clumps of up to 15 growths, each with 4–6

coriaceous, ligulate, dark green leaves 20–50 by 2–5 cm. Inflorescence arching or

suberect, with 1–7 flowers, green, sparsely papillose below. Pedicel and ovary glabrous,

4–7 cm long. Flowers 12–15 cm across. Dorsal sepal white with green venation at the

base, obovate, 3.8–4.3 by 2.2–2.5 cm. Synsepal pale green to white, broadly ovate,

3.5–3.8 by 2.3–2.6 cm. Petals light greenish, olive-green striped, with few black hairy

marginal warts near base, linear-tapering, strongly twisted, pendent, 8–11 by 1–1.1 cm,

with roundish hairy apex. Lip olive-green, deeply saccate, 4–4.5 by 2–2.5 cm. Staminode

obcordate, with a blunt basal umbo, three-lobed at apex, 9–11 by 6–7.5 mm, white

with dark green venation in the center. Fig. 21, h-k; 26, b.

Ecology. Coniferous, mixed and broad-leaved, evergreen, humid forests on highly

eroded, crystalline limestone at elev. 600–1450 m. Fl. September – November. Rare (EN).

Distribution. Vietnam (Cao Bang, Ha Giang, Hoa Binh, Lai Chau, Lao Cai,

Son La), S China.

Studied samples. Cao Bang, Bao Lac, CBL 394 (HN, LE); Ha Giang, Dong Van,

NTH 3364 (HN, LE), NTH 3539 (HN, LE); Hoa Binh, Mai Chau, HAL 680 (HN, LE), DKH 7871

(HN, LE, MO); Lai Chau, Averyanov s.n. (LE – photo); Lao Cai, Averyanov s.n. (LE – photo);

Son La, Moc Chau, DKH 7396 (HN, LE), NTH 2970 (HN, LE), HAL 9328 (HN, LE), HAL 9409

(HN); Son La, Yen Chau, DKH 7279 (HN, LE), HAL 9500 (HN, LE).

Notes. Species is closely allied to P. parishii from Myanmar, Thailand and

SW Yunnan, but differs in its lithophytic habit, glabrous inflorescences, bracts and

ovary and in white dorsal sepal.


Turczaninowia 2008, 11(1) : 5–168

Fig. 21. Paphiopedilum callosum var. callosum: a – flowering plant, b, c – staminode;

P. purpuratum: d – flowering plant, e–g – staminode; P. dianthum: h – flowering plant, i –

flower, j, k – staminode.

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Natural hybrids of Paphiopedilum species reported from Vietnam

P. appletonianum × P. gratrixianum. – P. × affine De Wild, 1906, Trib.

Hort. 1: 57.

Described from N Vietnam (“Tonkin”). Type (“cult. Bruxelles”) not located.

Any data on ecology and distribution are not yet available.

P. appletonianum × P. villosum. – P. × cribbii Aver. 2006, Orchids. Mag.

Amer. Orch. Soc. 76, 6: 458. Fig. 22, a, b; 26, c.

Described from S Vietnam (“Lam Dong Prov., Dalat City area…”). Type

(“November 14, 2005, L. Averyanov, P.K. Loc. HAL 8626”) – HN.

Ecology. Broad-leaved, evergreen and mixed forests on silicate soils at elev.

1500–2100 m. Fl. October – December. Very rare (DD).

Distribution. Vietnam (Lam Dong).

Studied samples. Lam Dong, Dalat area, HAL 8626 (LE - photo).

P. barbigerum (var. aspersum) × P. gratrixianum. Fig. 26, d.

Any data on ecology and distribution are not yet available. Fl. October.

Studied samples. Lam Dong, Dalat area, Averyanov s.n., 2005 (LE – photo).

P. callosum × P. gratrixianum. Fig. 26, e.

Any data on ecology and distribution are not yet available. Fl. October.

Studied samples. Lam Dong, Dalat area, Averyanov s.n., 2005 (LE – photo).

P. callosum × P. villosum. – P. × dalatense Aver. 2001, Orchid Digest, 65, 3:

134. – P. purpuratum auct. non (Lindl.) Stein: Seidenf. 1992, Orch. Indochina: 20;

Aver. 1994, Identif. Guide Vietnam. Orch.: 21. Fig. 22, c-e; 26, f.

Described from S Vietnam (“Lam Dong province, Don Duong…”). Type

(“cult. Dalat city Institute of Biology. Coll. N.V. Duy, No. P1 11, Nov. 1996”) – LE.

Ecology. Broad-leaved, evergreen and mixed forests on silicate soils at elev.

900–1700 m. Fl. April – October. Very rare (DD).

Distribution. Vietnam (Lam Dong).

Studied samples. Lam Dong, Don Duong, N.V. Duy, PI 11 (LE); Lam Dong, Dalat

area, HAL 4581 (LE – photo); Averyanov s.n., 2005 (LE – photo).

P. gratrixianum × P. villosum. Fig. 26, g.

Ecology. Broad-leaved, evergreen and mixed forests on silicate soils at elev.

900–2100 m. Fl. September – December. Rare (DD).

Distribution. Vietnam (Lam Dong).

Studied samples. Lam Dong, Dalat area, Averyanov s.n., 2005 (LE – photo).

P. helenae × P. hirsutissimum. – P. × herrmannii F. Fuchs et H. Reisinger,

1995, Linzer. Biol. Beitr. 27, 2: 1213. Fig. 22, f-i; 26, h.


Turczaninowia 2008, 11(1) : 5–168

Fig. 22. Paphiopedilum appletonianum × P. villosum (P. × cribbii): a – flowering plant, b – staminode;

P. callosum × P. villosum (P. × dalatense): c – flowering plant, d – flower, e – staminode;

P. helenae × P. hirsutissimum (P. × herrmannii): f – flowering plant, g – flower, h, i – staminode.

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Fig. 23. Paphiopedilum delenatii: a (Averyanov s.n., 2005); P. vietnamense: b (HAL 31); P. malipoense

var. malipoense: c (HAL 6207); P. malipoense var. jackii: d (HAL 169); P. malipoense

var. hiepii: e (Hiep s.n., 1995, type); P. micranthum: f (HAL 8341); P. hangianum: g (HAL

4789); P. emersonii: h (HAL 1581); P. concolor: i (HLF 5935).


Turczaninowia 2008, 11(1) : 5–168

Fig. 24. Paphiopedilum hirsutissimum var. esquirolei: a (Averyanov et al. s.n., 1995); P. hirsutissimum

var. chiwuanum: b (CBL 1588); P. barbigerum var. coccineum: c (P-10546); P. barbigerum

var. aspersum: d (P-10547); P. tranlienianum: e (HAL 183); P. helenae: f (Averyanov et al.,

CB 12, 1995, type); P. henryanum: g (NTH 3590); P. gratrixianum var. gratrixianum: h (Averyanov

s.n., 2005); P. gratrixianum var. daoense: i (Averyanov s.n., 2005).

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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 25. Paphiopedilum villosum var. villosum: a (Averyanov s.n., 2005); P. villosum var. annamense:

b (Averyanov s.n., 1995); P. villosum var. fusco-viride: c (Averyanov s.n., 2005); P. villosum

var. fusco-roseum: d (Averyanov s.n., 2005); P. villosum var. boxallii: e (Averyanov s.n., 2004);

P. appletonianum: f (HLF 5800); P. callosum var. callosum: g (HLF 5575); P. callosum var. warnerianum:

h (Averyanov, HAL s.n., 2004); P. callosum var. potentianum: i (Averyanov, HAL s.n., 2004).


Turczaninowia 2008, 11(1) : 5–168

Fig. 26. Paphiopedilum purpuratum: a (CBL 1458b); P. dianthum: b (Averyanov s.n.); P. appletonianum

× P. villosum: c (HAL 8626, type); P. barbigerum (var. aspersum) × P. gratrixianum:

d (Averyanov s.n., 2005); P. callosum × P. gratrixianum: e (Averyanov s.n., 2005); P. callosum ×

P. villosum: f (Averyanov s.n., 2005); P. gratrixianum × P. villosum: g (Averyanov s.n., 2005);

P. helenae × P. hirsutissimum: h (Averyanov s.n., 2002); P. malipoense × P. micranthum: i

(Averyanov s.n., 2006).

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Described from N Vietnam (“North Vietnam, border region to China”). Type

(“Herrmann 343/85”) – LI.

Ecology. Coniferous, mixed and broad-leaved, evergreen, closed forests on

limestone at elev. 700–900 m. Fl. August – October. Very rare (EN).

Distribution. Vietnam (Cao Bang).

Studied samples. Cao Bang, Trung Khanh, HAL 5627 (HN); Cao Bang, Tra Linh,

L. Averyanov et al. s.n., May 1997 (HN, LE).

P. malipoense × P. micranthum. – P. × fanaticum Koop. et N. Haseg. 1992,

Orch. Advocate, 18, 2: 50. Fig. 26, i.

Described from China (“ex China...”). Type (“… cult. H. Koopowitz, Paphanatics,

unlimited”) – K.

Ecology. Coniferous, mixed and broad-leaved, evergreen, closed forests on

limestone at elev. 800–1400 m. Fl. April – May. Very rare (DD).

Distribution. Vietnam (Cao Bang?, Ha Giang?).

Studied samples. Cultivated plants collected supposedly in highland limestone

areas of Cao Bang and Ha Giang provinces along border with China.

SUBFAMILY SPIRANTHOIDEAE DRESSLER

1979, Selbyana 5, 2 : 204.

Type: Spiranthes L.C. Rich

100–110(18) genera and 1200–1400(65) species. Tropical, subtropical,

temperate and boreal areas of all continents.

Trib. 3.1. Tropidieae Dressler

Dressler, 1983, Telopea 2, 4: 422.

Type: Tropidia Lindl.

2(2) genera and 30–40(3) species. Tropical and subtropical areas of all

continents.

Corymborkis Thouars, 1809, Nouv. Bull. Sci. Soc. Philom. (Paris) 2 ser. 1:

318; Rasmussen, 1977, Bot. Tidskr. 71, 3–4: 161–192.

Type: C. corymbis Thouars

Sympodial terrestrial herbs with short woody rhizome, fasciculate rigid roots

and erect semi-woody bamboo-like unbranched stems up to 4 m tall. Leaves sessile,

glabrous, plicate, sheathing at the base. Inflorescences lateral, panicle, with many

flowers. Flowers white, resupinate or not. Sepals and petals basally connate to form a

short tube, sub similar, long and narrow, widened toward the apex, spreading. Lip as

long as sepals, narrow and close to column, except apical ovate reflexed blade. Column

long, slender, straight, thickened to the apex. Anther erect, acute. Stigma broad, deeply

2-lobed. Pollinia 2, sectile, with linear caudicle and small peltate viscidium.

5(1) species. Tropical zone of the world.


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1. C. veratrifolia (Reinw.) Blume, 1859, Coll. Orch. Arch. Ind.: 125, pl. 43, 1,

pl. 42E; Seidenf. 1992, Orch. Indochina: 27; Aver. 1994, Identif. Guide. Vietnam.

Orch.: 29; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 24. – Hysteria

veratrifolia Reinw. 1828, Syll. Pl. Nov. Ratisbon. 2: 5.

Described from Java? Neotype (“Java: T. Lobb, 162” – sec. Rasmussen,

1977, Bot. Tidsskr. 71, 3–4: 170) – K.

Stem up to 5 m tall with spirally arranged leaves up to 45 cm long and 12 cm

wide, with long sheaths covering stem. Inflorescence up to 10 cm long. Sepals, petals

and lip 2–4 cm long. Reflexed apical blade of the lip ovate with finely crisped edges

and shortly pointed apex. Column to 2–3.5 cm long. Fruits cylindrical capsule 3–4 cm

long. Fig. 27, a,b; 31, a.

Ecology. All kinds of lowland and submontane forests on any soils. 50–1000 m.

Fl. May – July. Not rare (LR).

Distribution. Vietnam (probably all provinces). Tropical and subtropical Asia,

Australia and Pacific islands to Samoa.

Studied specimens. Bac Kan, Cho Don, HAL 4773 (HN, LE); Bac Kan, Na Ri, HAL

4952 (HN, LE), DKH 7631 (HN, LE, MO); Ha Nam Ninh, Cuc Phuong, Kien 147 CP (HN, LE);

Ha Son Binh, Luong Son, Tiep et al., NOT-2765 (HN, LE); Hai Phong, Cat Ba, 1988, LX-VN

3340, LX-VN 3779 (HN, LE); Hoang Lien Son, Dan Khao, 1964, sine coll. 3178 (HN, LE); Kim

Son, Trung, 1702 (HN, LE); Ninh Binh, Cuc Phuong national park, N.T. Hiep, NTH 5364 (HN);

Quang Binh, Bo Trach, HAL 6101 (HN, LE); Quang Binh, Ke Bang, VH 4609; VH 4768 (HN,

LE); Quang Binh, Minh Hoa, HAL 5870 (HN, LE), Lai Chau, Muong Te, HAL 10087 (HN, LE),

Nghe An, Tuong Duong, HLF 6606 (HN, LE).

Note. This tallest, shrub-like orchid in Vietnam is sometimes truly named as

“bamboo orchid”.

Tropidia Lindl., 1833, Bot. Reg. 19, tab. 1618; Seidenf. 1978, Dansk Bot.

Ark. 32, 2: 108–113. – Schoenomorphus Thorel ex Gagnep. 1933, Bull. Soc. Bot. Fr.

80: 351.

Type: T. curculigoides Lindl.

Sympodial terrestrial herbs with short woody rhizome, fasciculate rigid roots

and erect semi-woody rod-like branched, or superposed leafy stems. Leaves sessile,

glabrous, plicate, sheathing at the base. Inflorescences lateral or terminal, a short,

dense raceme. Flowers white, not resupinate. Sepals and petals free, or lateral sepals

connate into synsepalum. Lip short, spurred or saccate at the base, not mobile, narrowed

to down turned, triangular, obtuse apex. Column short. Clinandrium thin-margined.

Anther erect, dorsal. Pollinia 2, sectile, with linear caudicle and viscidium.

20(2) species. Tropical zone of the World.

Key to species

1. Leaves 2, broadly elliptic-ovate, usually sub opposite at a top of stem; inflorescence

terminal, more than 2 cm long; lateral sepals connate into synsepalum ...... 1. T. angulosa

– Numerous narrowly-ovate leaves distant throughout stem; inflorescence lateral, rare

terminal, less than 2 cm long; lateral sepals free or connate at the base only ... 2. T. curculigoides

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1. T. angulosa (Lindl.) Blume, 1858, Coll. Orch. Arch. Ind. Jap.: 122; Seidenf.

1992, Orch. Indochina: 27; Aver. 1994, Identif. Guide Vietnam. Orch.: 28; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 59. – Cnemidia angulosa Lindl.

1833, Bot. Reg. 19, tab. 1618.

Described from NE India (“India orientali; in montibus prov. Sylhet”). Lectotype

(“Wallich, 7388”) – K.

Stem to 40 cm tall, simple or superposed, with 2 apical sub-opposite leaves.

Leaves ovate to broadly-ovate, or even circular, acute to shortly acuminate, 6–20 cm

long, 2.5–11 cm wide. Inflortescence dense terminal spike 1–4 cm long on erect stalk

2–8 cm tall. Floral bracts narrowly-cuneate to filiform, 5–14 mm long. Flowers white,

sweet fragrant. Sepals oblong, 6–7 mm long, 3–4 mm wide, shortly suddenly apiculate

at the apex; lateral sepals connate into bidentate synsepalum. Petals smaller, oblong,

obtuse. Lip ovate, obtuse, 3.5–4.5 mm long, concave at the base, with strongly turned

down brightly yellow apex. Anther ovate, 2 mm long. Fig. 27, c; 31, b.

Ecology. All kinds of lowland and submontane forests on any soils. 0–1600 m.

Fl. April – July. Common (LR).

Distribution. Vietnam (probably all provinces). Tropical and subtropical mainland

Asia, S Japan, Taiwan, Hainan, Java, Sumatra.

Studied specimens. Bac Kan, Ba Be National Park, HLF 604 (HN, LE); Bac Kan,

Cho Don, HLF 879 (HN, LE); Cao Bang, Ba Be national park, Averyanov et al., CB 46 (HN, LE);

Cao Bang, Bao Lac, CBL 1547 (HN, LE); Cao Bang, Bao Lac, CBL 307 (HN, LE); Dak Lak, Chu

Yang Sin mt., HLF 5482 (HN, LE); Ha Tinh, Huong Son, HAL 1223, 5261 (HN, LE); Hai Phong,

Cat Ba, LX-VN 3266, 3335, 3396, 3447, 3660, 3679 (HN, LE, C); Hoa Binh, Lac Son, HAL

9033 (HN, LE); Hoa Binh, Pa Co, VH 2425 (HN, LE); Lai Chau, Sin Ho, HAL 9837 (HN, LE);

Nghe An, Tuong Duong, HLF 6625 (HN, LE), HLF 6708 (HN); Quang Binh, Ke Bang, VH

4606 (HN, LE); Quang Binh, Minh Hoa, HAL 5867 (HN, LE); Quang Ninh, Ha Long Bay, NTH

5621 (HN, LE); Son La, Mai Son, HAL 9586 (HN); Vungtau-Condao, Bay Canh, 1989, Averyanov

412 (HN, LE); Vungtau-Condao, Con Dao, 1987, Averyanov 881 (HN, LE); Vungtau-Condao,

Con Dao, 1989, Averyanov 512 (HN, LE).

2. T. curculigoides Lindl., 1840, Gen. Sp. Orch. Pl.: 497; Seidenf. 1992, Orch.

Indochina: 27; Aver. 1994, Identif. Guide Vietnam. Orch.: 27; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 60. – T. pedunculata Blume, 1859, Coll. Orch.

Arch. Ind.: 122, pl. 40; Seidenf. 1992, l.c.: 27. – T. graminea Blume, 1859, l.c.: 124, pl.

41, 3; Seidenf. 1992, l.c.: 27. – Schoenomorphus capitatus Thorel ex Gagnep. 1933,

Bull. Soc. Bot. France, 80: 351.

Described from NE India (“India orientali; montibus Sylhet, et secus ripas

fluminis Attran”). Lectotype (“Wallich, 7368”) – K.

Stem to 1.5 m tall, often branched, with numerous distant leaves. Leaves

narrowly-lanceolate to narrowly-ovate, acuminate, 5–25 cm long, 0.8–5 cm wide.

Inflorescence lateral axillary, rarely subapical, 1–1.8 cm long. Floral bracts triangularacuminate,

acute, 5–10 mm long, densely imbricate, lowest sterile. Flowers white.

Sepals and petals free, narrowly-ovate, acute, recurved, 7–10 mm long, petals smaller.


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Lip ovate, obtuse, 5–7 mm long, concave at the base, with turned down apex. Anther

narrowly-ovate, acuminate, 5–6 mm long. Fig. 27, d; 31, c.

Ecology. All kinds of forests on any soils. 0–2000 m. Fl. April – July. Very

common (LR).

Distribution. Vietnam (probably all provinces). Tropical and subtropical mainland

Asia, Taiwan, Hainan, Malaysia, Java, Kalimantan, Timor to New Caledonia.

Studied specimens. Bac Kan, Ba Be National Park, HLF 618 (HN, LE); Bac Kan,

Cho Don, Po Luong mt., HAL 4736 (HN, LE); Cao Bang, Ba Be national park, CB 43 (HN, LE);

Cao Bang, Tra Linh, CBL 1251 (HN, LE); Cao Bang, Yen Lac, CBL 307 (HN, LE); Ha Giang,

Quan Ba, NTH 3564 (HN, LE); Ha Son Binh, Luong Son, Tiep, NOT 2765 (HN, LE); Ha Tinh,

Huong Son, DKH 4381 (HN, LE, MO); Hai Phong, Cat Ba, LX-VN 3340a (HN, LE); Kon Tum,

Dak Gley, VH 1558 (HN, LE); Kon Tum, Dak Gley, VH 1383 (HN, LE); Lai Chau, Muong Te,

HAL 10094 (HN, LE); Lao Cai, Van Ban, DKH 6538, 6627 (HN, LE, MO); Nghe An, Tuong

Duong, HLF 6620 (HN, LE); Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4276 (HN, LE); Quang

Binh, Minh Hoa, HAL 5859 (HN, LE); Quang Ninh, Ba Mun, 1987, Averyanov 142 (HN, LE);

Quangnam-Danang, Phuoc Son, LX-VN 3088 (HN, LE); Thai Nguyen, Dong Hy, HAL 9093

(HN, LE); Thua Thien – Hue, A Luoi, HAL 7888 (HN, LE); Vungtau-Condao, Con Dao, 1987,

Averyanov 770, 806, 880 (HN, LE); Vungtau-Condao, Con Dao, 1989, Averyanov 535 (HN, LE).

Trib. 3.2. Cranichideae Endlicher

1842, Mant. Bot. Suppl. 2: 18.

Type: Cranichis Sw.

70–80(16) genera and 950–1050(62) species. Tropical and subtropical areas

of all continents, temperate and boreal regions of Northern Hemisphere.

Subtrib. 3.2.1. Goodyerinae Klotzsch

1846, in Hayne, Arzneygew 14, tab. 24 (sub “Goodyereae”).

Type: Goodyera R. Br.

38(15) genera and 475–500(61) species. Tropical and subtropical areas of all

continents, temperate and boreal regions of Northern Hemisphere, with highest diversity

in tropical Asia.

Anoectochilus Blume, 1825, Bijdr. 6, fig. 15; id. 1825, ibid. 8: 11 (sub “Anoecochilus”

– sphalm.); id. 1828, Fl. Javae 6, emend. nom. cons.

Type: A. setaceus Blume

Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome

rooting at nodes, ascending erect leafy floriferous stem and terminal lax, few flowered

spike. Leaves few, spirally arranged in rosette, petiolate, sheathing at the base, glabrous,

convolute, asymmetric, velvety dark green, brown-green to nearly black, with more or

less pronounced reticulation of white, yellow or pink nerves. Scape leafless, with 2–4

sterile yellowish bracts. Flowers, sessile, resupinate or not. Sepals free. Petals very

oblique, connivent to median sepal forming hood, lateral sepals spreading. Lip adnate

to column base, spurred. Spur conical to cylindric, with 2 short peltate glands on lateral

sides. Mesochile channeled, long to shortly fimbriate-toothed on flanges. Epichile with

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2 divergent flabellate spreading lobes. Column short, fleshy, with 2 wings at front, 2

lateral stigmas and prominent rostellum. Anther triangular-ovate, acute. Pollinia 2,

sectile, with stipe and viscidium.

30(7) species. Tropical mainland Asia, Indonesia, Australia, New Guinea, Pacific

Islands to Polynesia and adjacent islands.

Key to species

1. Spur and lip lie in one line or placed at a broad angle; spur more or less parallel with

ovary ...................................................................................................................................... 2

– Spur strongly bent down in right or acute angle with mesochile, placed in a wide

angle to ovary ..................................................................................................................... 3

2. Mesochile flanges long fimbriate; sepals flat without any callosities; epichile lobes

oblong-lanceolate, more or less flat ............................................................ 1. A. roxburghii

– Mesochile flanges toothed; sepals outside with few fat papillae along midvein;

epichile lobes triangular-flabellate, truncate, plicate along margin .......... 2. A. papillosus

3. Leaves black or gray-black, rarely dark gray-brown with green tint, commonly dense

reticulated with net of silvery-white nerves; mesochile flanges pectinate with more

than 10 small dents or short fringes ................................................................ 3. A. calcareus

– Leaves green, dark green to green-brown, rarely black-green, commonly reticulated

with net of white, yellowish or pink nerves; mesochile flanges with few blunt dents

on 2–6 long fringes ............................................................................................................. 4

4. Mesochile flanges with at least 1–2 long linear fringes 3–5 mm long ........................... 5

– Mesochile flanges with at few short dents ....................................................................... 6

5. Fringes on mesochile flange 4–6, all more or less equal in length; sepals lanceolate or

broadly-lanceolate, slightly falcate; column wings broad, rectangular, undulate along

margin ............................................................................................................ 4. A. albolineatus

– Mesochile with 1–3 pair of long fringes, other diminishing toward the base into small

narrow dents; sepals obovate, strongly oblique, with short suddenly attenuate apex;

column wings narrowly-triangular, straight, acute .................................. 5. A. annamensis

6. Spur narrowly-cylindrical, longer than 4 mm; epichile lobules lanceolate to broadlylanceolate,

4–5 times as long as broad; mesochile lobes insignificant, flanges narrow,

less than 1 mm wide, slightly erose or with few short irregular denticles near the base

of epichile ................................................................................................................. 6. A. lylei

– Spur broadly-conical shorter than 4 mm; epichile lobules narrowly obovate, commonly

not more than twice as long as broad; mesochile lobes large, erect, triangular, flanges

broad, triangular more than 1 mm wide, with 3–4 broad blunt dents along margin .........

............................................................................................................................ 7. A. brevilabris

1. A. roxburghii (Wall.) Lindl. 1840, Gen. Sp. Orch. Pl.: 499; Seidenf. 1992,

Opera Bot. 114: 32; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 11. –

Chrysobaphus roxburghii Wall. 1826, Tent. Fl. Napal.: 37. – A. setaceus auct. non

Blume: Aver. 1994, Identif. Guide Vietnam. Orch.: 46.

Described from N India (“India, Mt. Sylhet ...”). Syntypes (coll. F. De Silva

Wall. Cat. 7387; E India Company, icon. Wallich 34) – K.

Terrestrial herb with 3–4 petiolate leaves in basal rosette. Leaves ovate, acute,


Turczaninowia 2008, 11(1) : 5–168

to 5 cm long, velvety-green to dark olive-brown, with contrast reticulation of white,

yellow or pink nerves, rare only with 1–3 longitudinal colored nerves or uniformly

green. Scape pubescent, with 2–4 acuminate yellowish bracts, 12–20 cm tall, with 1–

15 lax flowers. Floral bracts ovate, acuminate, pubescent, 7–14 mm long. Ovary

glandular pubescent, 1–1.4 cm long. Sepals sub-similar, pink-brown, often with green

tint, ovate-oblong, 7–9 mm long, acute, glandular pubescent; median sepal with shortly

attenuate, recurved apex; lateral sepals at the base oblique. Petals white, often with

brownish-green tint along margin, oblique-falcate, as long as median sepal, attenuate.

Lip white, 1.4–1.8 cm long, spurred. Spur at a sharp angle to mesochile, 3–6 mm long,

conical, tapering to a minute 2-lobed apex, with 2 large peltate, warty glands. Mesochile

with 4–8 pairs of long fringes along flanges. Epichile lobes narrowly-obovate to oblong,

7–10 mm long. Column 3–4 mm tall, at front with 2 large semicircular parallel wings

not entering to the spur and 2 short rostellar arms. Anther ovate, acute, 3–4 mm long.

Capsule ellipsoid, 6–10 mm long, 6–7 mm wide. Fig. 27, e-m; 31, d, e.

Ecology. Closed broad-leaved forests of any kind, preferably on silicate soils.

200–1600 m. Fl. November – January, March. Not rare (VU).

Distribution. Vietnam (probably all provinces). Bhutan, Nepal, India, Sri Lanka,

S China, Thailand, Laos.

Studied specimens. Cao Bang, Tra Linh, VH 4966 (HN, LE); Cao Bang, Trung

Khanh, HAL 5562 (HN, LE); Dak Lac, Chu Yang Sinh mt., VH 6023 (HN, LE); Gia Lai Kontum,

Kontum, LX-VN 2204 (HN, LE); Gialai-Kontum, Kon Ha Nung, LX-VN 2858 (HN, LE); Ha

Giang, Quan Ba, HG 68 (HN, LE); Ha Giang, Yen Minh, NTH 3477 (HN, LE); Ha Tinh, Huong

Son, VA N.T. Hiep et al., 857 (HN, LE); Hoang Lien Son, Sapa, Takhtajan, 773 (LE); Khanh Hoa,

Khanh Le, VH 2482 (HN, LE); Khanh Hoa, Khanh Son, Hon Giao mt., VH 4271 (HN, LE); Gialai

Kontum, Kon Ha Nung, Kbang, LX-VN 2858 (HN, LE); Kon Tum, Dak Gley, VH 1943 (HN, LE),

VH 1944 (HN, LE), VH 1945 (HN, LE); Kon Tum, Kon Plong, DKH 4696 (HN, LE, MO);

Kontum, Dacuy, LX-VN 2204 (HN, LE); Kontum, Kon Plong, Mang La, VH 5069 (HN, LE); Lai

Chau, Sin Ho, HAL 9833 (HN, LE); Lam Dong, Lac Duong, Bi Dup mt., VH 3798 (HN, LE); Lam

Dong, Lac Duong, VH 4138 (HN, LE); Lang Son, Bac Son, HAL 6789 (HN, LE); Lao Cai, Van

Ban, HAL 2002 (HN, LE), HAL 2303 (HN, LE), HAL 2681 (HN, LE); Quangtri, Dongche,

Poilane; Son La, Thuan Chau, HAL 9735 (HN, LE); Tuyen Quang, Na Hang, HLF 690 (HN,

LE); Vanyen, Chapa, Petelot 12628 (P); Vinh Phu, Tam Dao, LX-VN 1356 (HN, LE), LX-VH

2382 (C, HN, LE).

Note. Widespread and very variable species, particularly in leaf coloration,

size of flower, length of spur and form of epichile lobes. Widely collected as a medicinal

plant used in traditional oriental medicine.

2. A. papillosus Aver. 2007, Taiwania, 52, 4: 287, fig. 1 & 2A.

Described from northern Vietnam (“Hoa Binh Prov., Mai Chau Distr., Pa Co

Municipality, Xa Linh village, Rung Gia forests”). Type (HAL 8978) – HN (holotype),

LE (isotype).

Terrestrial herb with 2–4 rosulate petiolate leaves. Leaves ovate to almost

circular, shortly acute, 2–4 cm long, velvety dark green, with reticulate network of

pink nerves. Scape 9–12 cm long, tomentose, with lax 3–5 flowers. Floral bracts,

97


98

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 27. Corymborkis veratrifolia: a – flower, b – sepal, petal, lip and column; Tropidia angulosa:

c – flower, side and frontal view; T. curculigoides: d – flower; Anoectochilus roxburghii: e –

flower and flattened column and tepals, f – spur section with glands; A. papillosus: g –

flowering plant and leaf, h, i – flower, frontal and side view, j – dorsal surface of median sepal

and petals, k – flattened tepals without lip, l – column, ventral and dorsal view, m – pollinarium.


Turczaninowia 2008, 11(1) : 5–168

triangular-cuneate, acuminate, 8–10 mm long, sparsely hairy. Ovary 10–12 mm long,

pubescent. Sepals light pink-brown, glabrous or sparsely hairy; dorsal sepal ovate, 6–

7 mm long, acuminate, with slightly upward reflexed apex; lateral sepals oblong-oblique,

obtuse, 7–8 mm long. Petals white, 6.5–7 mm long, 2–2.5 mm wide, strongly obliquefalcate,

rising from narrow base to oblique broad median part ending in attenuate

apex, each on dorsal surface with short longitudinal ridge bearing few massive white,

sometime forked capitate papillae. Lip white, with pink-brown tint at the spur apex,

13–14 mm long. Hypochile boat-shaped, basally with cylindric spur 3.5–4 mm long;

hypochile and spur, placed at narrow angle with mesochile; spur acutely bifid at apex,

with 2 large hemispheric sub-sessile glands inside. Mesochile 4–5 mm long, with narrow

base and 2 rectangular lateral lobes 2–2.5 mm wide, 3–3.5 mm long, denticulate with

5–10 irregular more or less low cuneate blunt dents. Epichile lobes spreading, obliquerectangular,

truncate, entire, 4.5–5.5 mm long, 2.5 mm wide. Column ovate, 4–4.5 mm

tall, with 2 short vertical wings at front; wings 1–1.4 mm long, about 1 mm tall, rising

at the middle part of column. Anther narrowly-ovate, acuminate, 3.5 mm long. Fig. 27,

g-m; 31, f.

Ecology. Broad-leaved humid evergreen forests on rocky limestone, commonly

in limestone pockets with deep soil. 900–1000 m. Fl. July – August. Rare (VU).

Distribution. Vietnam (Hoa Binh).

Note. Species is known up to now on alone type collection. From all known

representatives of the genus A. papillosus differs in petals bearing on dorsal surface

short longitudinal ridge ornamented with massive, often forked capitate papillae. On

other floral morphology this species approaches to Taiwanese A. koshunensis Hayata,

from which differs in large and broad, shallowly dentate side lobes of mesochile.

Obviously, this species is local endemic of rocky limestone remnant highlands spreading

in north-western Vietnam along Laotian border.

3. A. calcareus Aver. 1996, Bot. Journ. (St. Petersburg) 81, 10: 75, fig. 2.

Described from northern Vietnam (“Prov. Hoa Binh, near 1 km of Mai Chau

to the E”). Type (20 May 1995, Averyanov et al. s.n.) – LE.

Terrestrial and lithophytic herb with 3–4 rosulate petiolate leaves. Leaves ovate,

acute, to 7 cm long, dark velvety gray-brown to black, with dense network of silverywhite

to light pink nerves. Scape hairy, 8–12 cm long, with 3–7 lax flowers. Floral

bracts cuneate, tomentose, as long as ovary. Ovary tomentose, 8–10 mm long. Sepals

light green, sparsely hairy; dorsal sepal broadly-ovate, 4–5 mm long, acute; lateral

sepals ovate, oblique, 6–7 mm long. Petals white, greenish to the apex, strongly falcate,

attenuate to the base and apex, as long as median sepal. Lip white, 13–15 mm long,

basally with small conical, flattened spur in a wide angle to a claw; spur 3 mm wide,

2.5 mm wide, with 2 capitate gland. Claw 4–5 mm long, with 10–15 pair of dents or

short fringes on flanges. Epichile lobes obovate, 5–6 mm long 2.5–3 mm wide. Column

3 mm tall, with 2 small vertical wings broadening proximally, not entering into spur.

Anther ovate, acute. Fig. 28, a-e; 31, g.

99


100

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Ecology. Coniferous and broad-leaved forests on rocky limestone, commonly in

limestone pockets with deep soil. 450–1600 m. Fl. January – February. Occasional (VU).

Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Hoa Binh, Lai Chau,

Ninh Binh, Quang Binh, Son La, Thanh Hoa).

Studied specimens. Bac Kan, Cho Don, HLF 803 (HN, LE), HAL 4735 (HN, LE);

Cao Bang, Nguyen Binh, CBL 1583 (HN, LE); Ha Giang, Bac Me, HAL 6485 (HN); Ha Giang,

Quan Ba, NTH 3573 (HN, LE), HG 74 (HN, LE); Hoa Binh, Mai Chau, HAL 843 (HN), DKH

7801 (HN, LE, MO), DKH 8028 (HN, LE, MO); Lai Chau, Sin Ho, HAL 10058 (HN); Ninh Binh,

Cuc Phuong national park, HAL 1696 (HN); Quang Binh, Bo Trach, HAL 6055 (HN); Quang

Binh, Minh Hoa, Ke Bang, VH 4764 (HN, LE), HAL 5984 (HN), HAL 5866 (HN, LE – photo);

Son La, Moc Chau, DKH 7437 (JM. LE, MO), HAL 9305 (HN); Thanh Hoa, Ba Thuoc, HAL

2994 (HN, LE – photo), HAL 3439 (HN); Thanh Hoa, Quan Hoa, HAL 3742 (HN), Son La, Yen

Chau, HAL 9512 (HN, LE).

Note. Typical herbaceous element of rocky limestone at the north of Vietnam.

4. A. albolineatus Par. et Reichenb. f. 1874, Trans. Linn. Soc. Bot. 30: 141. –

A. siamensis Schleichter, 1906, Feddes Repert. 2: 83; Seidenf. 1992, Opera Bot. 114:

32; Aver. 1994, Identif. Guide Vietnam. Orch.: 46; Aver. et Averyanova, 2003, Updated

Checklist Orch. Viet.: 11.

Described from Myanmar (“Burma – Moulmein”). Type (“Parish 325”) – K

(holotype), W (isotype).

Terrestrial herb with 2–3 shortly petiolate leaves in basal rosette. Leaves

broadly–ovate, acute, 1.5–4 cm long, velvety dark-green to brown-green or nearly

black, often with broad irregular median longitudinal white to yellowish stripe or

reticulation of pink to yellowish veins. Scape pubescent, with 2–3 acuminate yellowish

bracts, 6–15 cm tall, with 1–8 lax flowers. Floral bracts ovate, acuminate, pubescent.

Ovary pubescent, 5–10 mm long. Sepals sub-similar, ovate-oblong, 7–10 mm long,

acute, sparsely hairy. Petals narrowly-obovate, slightly oblique, as long as median

sepal, shortly acuminate. Lip 5–7 mm long, spurred. Spur directed parallel with ovary

and claw, 1.5–3 mm long, broadly-conical, notched at the apex, with 2 large peltate,

warty glands. Mesochile with 4–7 pairs of long fringes along flanges. Epichile lobes

narrowly-obovate to oblique-flabellate, more or less truncate, 5–7 mm long. Column at

front with 2 large narrowly-triangular, parallel wings not entering to the spur and 2 short

broad rostellar arms. Anther narrowly-ovate, acute, 3–4 mm long. Fig. 28, f-h; 31, h.

Ecology. Humid broad-leaved forests on silicate soils. 1100–1700 m. Fl. November

– January. Rare (VU).

Distribution. Vietnam (Kon Tum, Lam Dong, Lao Cai), Myanmar, Thailand.

Studied specimens. Kon Tum, Dak Gley, VH 1675 (HN, LE); Lam Dong, Lac

Duong, Hon Giao ridge, VH 4132 (HN, LE); Muang Xen, Hayata s.n. (TI).

5. A. annamensis Aver. 2005, Rheedea, 15, 2: 83, fig. 1 & 4a, b.

Described from central Vietnam (“Thua Thien – Hue Prov., Huong Thuy

Distr., Duong Hoa Municipality, Huong Thuy forest enterprise ... Mang Chan ridge”).


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101

Fig. 28. Anoectochilus calcareus: a – flowering plant, b – flower, c – flattened tepals, d – flattened

lip and section of spur, e – column, dorsal and ventral view; A. albolineatus: f, g – flattened

tepals and lip, h – column ventral and side view; A. annamensis: i – flowering plant, j – flattened

tepals, k – flattened lip and section of spur, l – column and section of spur, side view;

A.lylei: m, n – flattened tepals and lip, o – column and spur section, frontal view, p – column, side view.


102

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Type (HAL 8177) – HN (holotype), LE (isotype). d-EXSICCATES OF VIETNAMESE

FLORA 0023/HAL8177.

Terrestrial herb with 2–3 rosulate, shortly petiolate leaves. Leaves ovate, finely

undulate along margin, shortly acute, 3–5 cm long, dark velvety green, olive-green to

nearly black, with reticulate network of pink nerves. Scape 7–16 cm long, tomentose,

with 3–10 lax flowers. Floral bracts triangular-cuneate, acuminate, 4–6 mm long.

Ovary 6–10 mm long, densely pubescent with simple and glandular hairs. Sepals dark

olive-brown to reddish-brown, pubescent; dorsal sepal ovate, 4.5 mm long, acuminate,

with strongly upward reflexed apex; lateral sepals oblong, obtuse, with broad asymmetric

base, 7–9 mm long. Petals light green, 4.5 mm long, 3–3.2 mm wide, strongly obliquefalcate,

rising from very narrow base to oblique broad apical part ending in attenuate

tip. Lip white with light green fringes, about 17 mm long. Hypochile boat-shaped,

basally with broadly-conical spur, 3 mm long, 2 mm broad; spur not bent, placed in one

line with lip, notched at apex, with 2 large flat stalked semicircular glands inside.

Mesochile form narrow claw 1.5 mm long, with fleshy upturned margins and 2–5

back reflexed lateral dents or fringes 1–3.5 mm long. Epichile lobes spreading, oblong

to oblong-lanceolate, 8 mm long, 3 mm wide. Column 5–5.5 mm tall, with 2 vertical

wings at front; wings 2–2.5 mm long, cuneate or narrowly-triangular, acute, rising

from the middle of the column. Anther large, narrowly-ovate, about 4 mm long. Fig.

28, i-l; 31, i; 32, a.

Ecology. Humid broad-leaved evergreen forests on silicate soils. 100–1000 m.

Fl. March – May. Occasional (VU).

Distribution. Vietnam (Thua Thien – Hue, Vinh Phuc).

Studied specimens. Tam Dao, LX-VN 874 (LE); Thua Thien – Hue, A Luoi, HAL 7172

(HN), HAL 7271 (HN; LE), HAL 7621 (HN; LE), HAL 7766 (HN); Thua Thien – Hue, Nam Dong,

HAL 6897 (HN; LE); Thua Thien - Hue, Xuan Loc, HLF 1667 (HN, LE), HLF 1825 (HN, LE).

Note. Species superficially resemble widespread and common species A. roxburghii,

but actually belongs to group of species with straight (not bent) short spur like

A. brevilabris and A. albolineatus. This often overlooked species may be widespread

and quite common in low mountain areas of Vietnam.

6. A. lylei Rolfe ex Downie, 1925, Kew Bull. 1925: 411; Seidenf. 1992, Opera

Bot. 114: 33; Aver. 1994, Identif. Guide Vietnam. Orch.: 44; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 11.

Described from Thailand (“Siаm ... Me Awn, 750 m”). Type (“Lyle in herb.

Kerr, 297”) – K.

Terrestrial herb with 2–4 petiolate leaves in basal rosette. Leaves ovate to

almost circular, 2–6 cm long, shortly acute, dark green, green-brown to nearly black

with reticulated yellowish to pink nerves, rarely with only 1 or 3 longitudinal colored

veins. Scape densely pubescent, with 2–4 yellowish, broad, acuminate bracts, 6–15 cm

tall, with 1–8 lax flowers. Floral bracts ovate, acuminate, pubescent, 7–14 mm long.

Ovary 10–16 mm long, densely pubescent. Sepals sub-similar, ovate-oblong, 7–11 mm

long, shortly acute, hairy. Petals obovate, as long as median sepal, strongly oblique,


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103

rising from narrow base to broad median part and oblique attenuate apex. Lip 15–

18 mm long, spurred. Spur cylindric, directed parallel with ovary and claw, rare at

wide angle with claw, 4–5 mm long, with 2 large peltate, warty glands. Mesochile

boat-shaped, with short claw, along flanges slightly erose or with few short irregular

denticles. Epichile lobes lanceolate to broadly-lanceolate, 8–10 mm long, 1.2–1.8 mm

wide. Column 3–4 mm tall, at front with 2 narrowly-triangular, acute wings, not entering

to the spur and short bifid rostellum. Anther ovate, acute, 4 mm long. Fig. 28, m-p; 32, b.

Ecology. Closed humid broad-leaved forests on silicate soils. 600–2000 m. Fl.

November – January. Rare (VU).

Distribution. Vietnam (Dak Nong, Gia Lai, Khanh Hoa, Kon Tum, Lam Dong,

Quang Tri), Thailand.

Studied specimens. Da Tan La river, Col de Pren, Tixier drawing (P); Dak Nong,

Dak Glong, Ta Dung mt., HLF 5656 (HN, LE); Dalat, Evrard 1944, 2029 (P); Gialai-Kontum,

1985, Averyanov et al., LX-VN s.n. (HN, LE); Khanh Hoa, Khanh Son, Hon Giao ridge, VH 4276

(HN, LE); Kon Tum, Dak Gley, VH 2278a (HN, LE); Kontum, Kon Plong, Mang La, VH 5599a

(HN, LE); Kontum, Sa Tay, Chu Mom Ray, P-7906a (HN, LE); Langbian, Rolfe; Phu Sapoum,

Blao, Poilane 23804 (P); Quangtri, Poilane 11288 (P).

7. A. brevilabris Lindl. 1840, Gen. Sp. Orchid. Pl.: 499. – A. griffithii Hook.

f. 1890, Fl. Brit. Ind. 6: 96. – A. sikkimensis King et Pantl. 1896, Journ. As. Soc.

Bengal, 65: 124; id. 1898, Ann. Bot. Gard. (Cаlcuttа) 8, tab. 391; Aver. 1994, Identif.

Guide Vietnam. Orch.: 46; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.:

11. – A. chapaensis Gagnep. 1931, Bull. Mus. Hist. Nat. (Paris) 2 ser. 3, 7: 679;

Seidenf. 1992, Opera Bot. 114: 33; Aver. 1994, Identif. Guide Vietnam. Orch.: 44;

Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 10. – A. tridentatus Seidenf.

1992, Opera Bot. 114: 33, fig. 11; Aver. 1994, Identif. Guide Vietnam. Orch.: 44; Aver.

et Averyanova, 2003, Updated Checklist Orch. Viet.: 11.

Described from NE India (“India, Assam”). Type (“Mack s.n.”) – K.

Terrestrial herb with 3–4 petiolate leaves in basal rosette. Leaves dark green,

with yellow to pink nerve reticulation, ovate, to broadly-ovate, shortly acute, to 6 cm

long. Scape pubescent, to 15 cm long, with 1–6 lax flowers and 2–3 acuminate sterile

bracts. Floral bracts ovate, acuminate, 6–10 mm long. Sepals sub-similar, olive-green,

oblong, obtuse, 6–8 mm long. Petals oblique-falcate, as long as median sepal. Lip

white, spurred, 8–10 mm long; spur short, 3–4 mm long, broadly-conical, bifid at the

apex, with 2 rather small peltate glands. Mesochile with 3–4 pairs of short blunt teeth

on flanges. Epichile lobes divergent, flabellate-triangular, truncate, 7–8 mm long. Column

3–4 mm tall, with 2 narrowly-triangular, obtuse, parallel wings at front and broad bifid

acute rostellum. Fig. 29, a, b.

Ecology. Humid broad-leaved forests on silicate soils. 900–1500 m. Fl.

December – February. Rare (EN).

Distribution. Vietnam (Lao Cai, Vinh Phuc), Bhutan, NE India.

Studied specimens. Tamdao, Hayata s.n. (TI); Tonkin: Cha-pа, Petelot 5147 (P);

Vinh Phu, Tam Dao, LX-VN 874 (HN, LE).


104

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Note. Some specimens from northern Vietnam have intermediate morphology

between this species and A. annamensis.

Cheirostylis Blume, 1825, Bijdr. 6, fig. 16; id. 1825, ibid. 8: 413.

Type: C. montana Blume

Small terrestrial, lithophytic and epiphytic sympodial herbs. Rhizome creeping,

with very swollen, succulent internodes and roots modified into ridges or pillows densely

covered by root hairs. Stem ascending, leafy at the base, with erect scape and lax

terminal spike. Leaves few, spirally arranged, petiolate or sessile, sheathing at the

base, glabrous, convolute, usually dull green, acute. Scape leafless, with 1–5 sterile

hyaline bracts. Flowers small, sessile, resupinate, hardly opening. Sepals sub-similar,

connate for half of their length to form a swollen tube. Petals connivent or joined to

median sepal forming hood. Lip spurless. Hypochile concave, with various papillae on

lateral sides, or lacking any glands. Epichile reflexed, 2-lobed, rarely entire, connected

to hypochile by entire simple short claw. Column short, erect, with 2 thin prominent

stelidia and 2 rostellar arms. Anther triangular-ovate, acute. Pollinia 2, sectile, with

stipe and viscidium. Stigma with 2, small, convex, separated lobes.

50(13) species. Subtropical and tropical zone of the Eastern Hemisphere with

main center of diversity in mainland SE Asia.

Key to species

1. Leafy stem creeping; leaves distant, distichous, sessile, 4–7 mm long ....... 1. C. serpens

– Leafy stem ascending to erect; leaves rosulate, spirally arranged, petiolate, usually

longer than 7 mm ................................................................................................................. 2

2. Inflorescence sub-sessile, scape less than 1 cm long, much shorter than apical leaves

................................................................................................................................... 2. C. inabai

– Leafy stem ascending to erect; leaves rosulate, spirally arranged, petiolate, usually

longer than 7 mm; scape taller than 2 cm ........................................................................... 3

3. Stem erect; leaves distant in lower part of stem, erect to sub-erect, to 6 cm long,

cuneate at the base, sub-succulent, brightly uniform green, glossy; stigma lobes

cylindric-conical ..................................................................................................... 3. C. foliosa

– Stem ascending; leaves usually rosulate at the base of stem, spreading, less than

4cm long, broadly-triangular to cordate at the base, thin, velvety dull-green to dark

green, commonly with dark veins; stigma lobes hemispheric ........................................ 4

4. Epichile narrow, simple, ligulate-oblong, longer than broad; hypochile at the base

without glands or with hardly visible rudimentary warts ................................ 4. C. takeoi

– Epichile distinctly broader than long, often 2-lobed, with toothed or fringed lobes;

hypochile at the base with slender finger-like glands placed along lateral veins ....... 5

5. Lobes of epichile deeply toothed of fringed along margin ............................................. 6

– Lobes of epichile entire, finely irregular denticulate or shallowly crenulate ............... 10

6. Scape 6–9 cm long with 1–2 flowers .................................................................................. 7

– Scape normally longer than 9 cm, with 1–10 flowers ....................................................... 8

7. Leaves glossy, more or less rigid, silvery-gray to light green, with network of dark

green nerves; petals narrowly-obovate to oblanceolate; epichile lobes with 5–8


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105

triangular dents; hypochile with numerous fat conic glands along lateral veins .................

....................................................................................................................... 5. C. marmorifolia

– Leaves uniform dull gray-green, thin and soft; petals filiform-linear; epichile lobes

with 4 finger-like fringes; hypochile with 2 slender glands along lateral veins .....................

............................................................................................................................... 6. C. filipetala

8. Sepals white, 3.5–5 mm long; epichile 1.5–3.5 mm long and wide; epichile lobes usually

with 4–7 dents; hypochile on sides with 2(3) glands ..................................... 7. C. chinensis

– Sepals light olive-green to greenish-pink, 4.5-6.5 mm long; epichile 4–7 mm long and

wide; epichile lobes usually with 7–12 dents or fringes; hypochile on sided with 3–5

or more glands ..................................................................................................................... 9

9. Epichile lobes deep lacinate, with long finger-like fringes; leaves usually dark green

with light green median stripe .............................................................. 8. C. cochinchinensis

– Epichile lobes deeply dentate, with narrowly-triangular teeth; leaves usually light

dull gray-green with dark veins .................................................................. 9. C. yunnanensis

10. Scape less than 5 cm tall; lip 5–5.5 mm long; epichile entire, transverse-rhomboid ........

..................................................................................................................... 10. C. thanmoiensis

– Scape normally taller than 5 cm; lip longer than 6 mm; epichile distinctly emarginate or

2-lobed ................................................................................................................................... 11

11. Epichile 2–3 mm wide, emarginate to 2-lobed, with entire semi-circular lobes .....................

.......................................................................................................................... 11. C. bipunctata

– Epichile more than 5 mm wide, distinctly 2-lobed, with flabellate to oblique-rectangular

lobes, crenulate or denticulate along margin ................................................................. 12

11. Epichile lobes spreading, to 4 mm long, shallowly crenulate or broadly dentate, with

2–4(5) blunt-roundish indistinct dents; petals obovate, much broadening to the apex

............................................................................................................................ 12. C. latipetala

– Epichile lobes convergent, 10–14 mm long, irregular denticulate, with low irregular rounded

teeth; petals elliptic, narrowing to the base and apex .............................. 13. C. spathulata

1. C. serpens Aver. 2005, Rheedea, 15, 2: 87, fig. 5 & 8a, b.

Described from northern Vietnam (“Quang Binh Prov., Bo Trach Distr., Phong

Nha – Ke Bang national park ...”). Type (“24 January 2005, L. Averyanov, P.K. Loc,

N.T. Vinh, HAL 6092)” – HN (holotype), LE (isotype).

Dwarf creeping epiphyte or lithophyte with succulent green stem densely

adpressed to the ground. Stem with many distichous, sessile, cordate, acute or apiculate,

brightly emerald-green leaves, 4–7 mm long, 4–5 mm wide. Scape leafless, ascending,

1.5–1.8 cm long, sparsely haired, with 2–3 light greenish, ovate, apiculate bracts, and

1–3 flowers. Sepals green, obtuse, sparsely haired, 3.5 mm long, connate on half way

to tip from base; dorsal sepal ovate, about 2.5 mm wide, lateral sepals narrowly-ovate

to oblong, about 1.5 mm wide. Petals white, spathulate, falcate, broadening from very

narrow base to rounded apex, 3–3.5 mm long, 1–1.4 mm wide. Lip white about 2.5–

3.5 mm long. Hypochile concave, with central swelling and 2 slender, rather straight

gland on each side. Epichile white, with 2 light olive-green to yellow oblong spots at

the center, 4 mm wide, 2-lobed; lobes squarish, entire, slightly plicate. Column 2 mm

tall, with large, narrowly-ovate anther; rostellum arms subulate, acute, as long as or

little longer than anther; stelidia large, of the same length, spathulate, with obtuse to


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

roundish apex. Fig. 29, c-e; 32, c.

Ecology. Wet broad-leaved forests on rocky limestone. 600–900 m. Fl. January–

February. Very rare (EN).

Distribution. Vietnam (Bac Kan, Quang Binh).

Studied specimens. Bac Kan, Cho Don, HAL 4804 (HN, LE).

2. C. inabai Hayata, 1914, Ic. Pl. Formos. 4: 108, fig. 56. – C. pingbianensis

K.L. Lang, 1996, Acta Phytotax. Sin. 34, 6: 635. – C. phamhoangii N.T. Tich, 2001,

Orch. Vietnam, 1: 197, nom. inval.

Described from Taiwan (“Rinkoho, ad 5000 ped. alt.”). Type (“H. Inaba,

Aug., 1911”) – TI?

Terrestrial creeping herb with ascending fleshy pink stem 10–15 cm tall. Leaves

distant and small at the base, clustered at the apex with 4–5 narrowly-ovate, attenuate,

petiolate, green leaves, 3–4 cm long, 1.5–2 cm wide. Petiole 1–1.5 cm long. Inflorescence

very short, 1–2 cm long, secund, with 3–7 flowers, clustered in dense headlike

spike. Ovary narrowly-obovoid, to 1 cm long, glabrous. Flowers pure white. Sepals

sub-similar, forming cylindric tube 1 cm long, with 3 teeth at apex. Petals broadlylanceolate,

1 cm long, 3 mm wide. Lip 1.3 cm long. Hypochile saccate, boat-shaped,

at the base with few small close glands along each lateral vein. Epichile conduplicate,

broad, 2-lobed, 6–8 mm wide; lobes spreading; each lobe dissected into 5 close fingerlike

segments 3–4 mm long, 0.5–0.6 mm wide. Column short. Stelidia narrow-parallel,

4 mm long, sub-equal in length with rostellum arms. Anther broafly-ovate, much shorter

than rostellum. Fig. 29, f-i.

Ecology. Ericaceous thickets on silicate soils along tops of ridges. 2000–2150m.

Fl. September. Very rare (EN).

Distribution. Vietnam (Lam Dong), S China, Taiwan.

Studied specimens. Lam Dong, Lac Duong, Langbian mt., T.07.09.93a (icon

“Cheirostylis phamhoangii”, N.T. Tich, 2001, l.c.: 199).

Note. Very rare species known in Vietnam by the only collection.

3. C. foliosa Aver. 2008, Taiwania, 52, 4: 295, fig. 2e & 7.

Described from northern Vietnam (“Son La Prov., Moc Chau Distr., Chieng

Hac Municipality”). Type (“31 Oct. 2006, N.T. Hiep, L. Averyanov, P.V. The, HAL

9415”) – HN (holotype), LE (isotype).

Erect lithophytic herb. Stem 10–16 cm tall, leafy and glabrous in lower third.

Leaves 2–4, sessile or sub-sessile, narrowly-ovate, erect, acute, 2.5–6 cm long, pure

light green, sub-succulent, glossy. Scape 8–14 cm long, densely pubescent in upper

part. Rachis densely pubescent, 1.5–2.5 cm long with 5–8 flowers. Floral bracts

broadly-ovate, acuminate, sparsely glandular ciliate along margin, 5–6 mm long. Ovary,

narrowly-ovate, 4–4.5 mm long, densely pubescent with glandular hairs. Sepals olivegreen

to dull pink-brown, sub-similar, ovate, 2.8–3.2 mm long, connate on 2 / 3 from the

base, sparsely pubescent. Petals white, spathulate, falcate, broadening from narrow

base to rhomboid rounded apex, slightly shorter than sepals. Lip white, 3.5 mm long


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107

and wide. Hypochile concave, cup-like, 1.5 mm across, with 1 large longitudinal cristate

straight ridge along lateral veins. Epichile papillose, 1.2 mm wide, 2-lobed, lobes square,

entire or with 2 indistinct rounded dents. Column 2 mm long, 1.4 mm wide; anther cup

lanceolate; rostellum arms narrowly spathulate, slightly shorter than anther cup; stelidia

shorter, thick, cylindrical, obtuse, with papillose surface. Fig. 29, j-o; 32, d.

Ecology. Broad-leaved forests on rocky limestone, commonly on mountain

tops. 1200–1300 m. Fl. September – November. Very rare (EN).

Distribution. Vietnam (Son La).

Note. Very rare species known by the only type collection.

4. C. takeoi (Hayata) Schleichter, 1919, Fedde Repert. Beih. 4: 171; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 20. – Arisanorchis takeoi Hayata,

1914, Ic. Pl. Formos. 4: 110, fig. 57. – C. eglandulosa Aver. 1996, Bot. Journ. (St.

Petersburg), 81, 10: 80, fig. 5.

Described from Taiwan (“Arisan: inter Karapin et Suisharyo, ad 3300 ped.

alt.”). Type (“Takeo Ito, Mart. 28, 1914”) – TI?

Lithophytic herb 5–16 cm tall with 1–3 basal, shortly petiolate leaves. Leaf

blade dull silvery-green with unclear green vein network, ovate to broadly-ovate, to

3.5 cm long. Scape and rachis pubescent, rachis 0.5–1 cm long, with 1–5 lax, hardly

opening flowers. Floral bracts triangular-ovate, acuminate, 3–8 mm long, not longer

than ovary, dense tomentose. Ovary obovoid, 3–7 mm long, densely pubescent. Sepals

outside sparsely pubescent, light olive-pink, narrowly-ovate, 5–7 mm long, joined on 2 / 3

of their length into a tube. Petals white, free, falcate-lanceolate, 5–6.5 mm long, 1–1.5

mm wide. Lip white, 6–7 mm long. Hypochile concave, having no glands or with

hardly visible rudimentary papillae inside. Epichile strongly bended down, narrowlyovate

to oblong-lanceolate, entire, obtuse at the apex. Column 1–1.5 mm tall. Fig. 29,

p-s; 32, e.

Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.

600–1000 m. Fl. March – April. Not rare (VU).

Distribution. Vietnam (Cao Bang, Ha Giang, Lang Son), Ryukyu Islands,

Taiwan.

Studied specimens. Cao Bang, Tra Linh, Averyanov et al., CB32a, 22 Oct.

1995 (LE); Cao Bang, Bao Lac, CBL 1372 (HN, LE); Cao Bang, Trach An, 5 Dec.

1998, Averyanov et al. s.n. (HN, LE); Ha Giang, Dong Van, CBL 1760b (HN, LE);

Ha Giang, Quang Ba, DKH 4895 (HN, LE, MO); Lang Son, Bac Son, HAL 6708

(HN, LE), HAL 6709a (HN, LE).

Note. Typical element of herbaceous lithophytic vegetation on mossy rocky

solid limestone.

5. C. marmorifolia Aver. 2000, Turczaninowia, 3, 2: 19. – C. marmorata

Aver. 1999, Lindleyana, 14, 4: 216, non Lindl. ex Lem., 1848.

Described from northern Vietnam (“Cao Bang Province, Ha Lang District,

Thanh Nhat Municipality”). Type (“27 November 1998, L. Averyanov, P.K. Loc,

N.X. Tam, CBL s.n.”) – LE.


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Fig. 29. Anoectochilus brevilabris: a – flattened tepals and lip, b – column, frontal and side view;

Cheirostylis serpens: c – flowering plant, d – flattened tepals, e – column, side view; C. inabai:

f – flowering plant, g – flowers, h – flattened sepals, petal and lip, i – column and lip, side view;

C. foliosa: j – flowering plants, k – flower, l, m – flattened tepals and lip, n – column, dorsal,

frontal and side views, o – floral bract; C. takeoi: p – flowering plant and leaf, q – flower, r –

flattened tepals, s – lip, side view.


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109

Dwarf lithophytic herb with creeping chain rhizome and ascending short stem

to 6 mm tall with 2–4 rosulate leaves. Leaves shortly petiolate, ovate, rigid, to 2 cm

long, light green to silvery white with contrasting dark green veins. Scape erect,

pubescent, 6–8 cm tall, with 1–2 flowers. Floral bracts pubescent, 3–4 mm long, shorter

than ovary. Ovary densely pubescent, 4 mm long. Sepals olive-green, pubescent, ovate,

5 mm long, joined on 2 / 3 of their length forming tube. Petals white, oblanceolate, falcate,

5.5 mm long. Lip 8 mm long, white, with 2 green spots at the base of epichile. Hypochile

concave, with fleshy central keel and numerous fleshy glands on each side. Epichile

2-lobed, lobes deeply fringed, with 6–7 cuneate teeth. Column 2.5 mm tall. Fruit ovoid

capsule 4 mm long. Fig. 30, a-e; 32, f.

Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.

400–1500 m. Fl. December – March. Rare (EN).

Distribution. Vietnam (Bac Kan, Cao Bang, Son La).

Studied specimens. Bac Kan, Ba Be national park, HLF 508 (HN, LE), HLF 650

(HN, LE); Cao Bang, Bao Lac, CBL 1485 (HN, LE); Cao Bang, Tra Linh, 27 May 1997,

L.Averyanov, N.T. Hiep, VH s.n. (LE); Son La, Yen Chau, DKH 7203 (HN, LE).

6. C. filipetala Aver. 2003, in Aver. et Averyanova, Updated Checklist Orch.

Viet.: 20, 76, fig. 4.

Described from southern Vietnam (“Kon Tum Prov., Sa Thay Distr., Municipality

Sa Son, Chu Mom mountain ...”). Type (“17 January 2000, P.K. Loc et al., P-

8498”) – HN (holotype), LE (isotype).

Epiphytic herb with ascending leafy stem to 4 cm long and 3–5 distant petiolate

sheathing leaves. Leaf blade dull green, triangular-ovate to cordate, acute, to 1.1 cm

long, 0.8 cm wide. Scape 5–9 cm tall, densely pubescent, with 1–2 flowers. Floral

bracts triangular, 3–5 mm long, much shorter than ovary. Ovary 8–10 mm long, glabrous.

Sepals yellowish-white, sparsely hairy, 4 mm long, dorsal sepal broadly-ovate, lateral

sepals narrowly-ovate acute. Petals white 4 mm long, filiform, slightly broadening to

the apex. Lip white, with 2 light green spots at the epichile base, 6 mm long. Hypochile

concave, boat-shaped, at the base with 2 slender, finger-like glands along lateral veins.

Epichile 2-lobed, lobes fringed, with 4 finger-like segments each. Column narrow,

3mm long, with 2 long stelidia and rostellum arms. Anther narrowly-ovate, acite,

1.5 mm long. Fruit narrowly-ellipsoid, 8–11 mm long. Fig. 30, f-i.

Ecology. Primary wet broad-leaved closed forests on granite. 900–1000 m.

Fl. January. Very rare (DD).

Distribution. Vietnam (Kon Tum).

Note. Very rare species known from the only type collection.

7. C. chinensis Rolfe, 1895, in Hemsl. Ann. Bot. (London) 9: 158; Seidenf.

1992, Opera Bot. 114: 35; Aver. 1994, Identif. Guide Vietnam. Orch.: 41; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 20.

Described from Taiwan and Hongkong (“Formosа ... ; Hongkong”). Lectotype

(“Henry, 320”) – K.


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Terrestrial and lithophytic herb. Stem 6–30 cm tall, with 1–5 leaves at the

base. Leaves dull gray-green, narrowly-ovate to ovate-cordate, acute, 1–3 cm long.

Scape tomentose, to 22 cm long. Rachis to 1.5 cm long, tomentose, with 1–10 flowers.

Floral bracts triangular-ovate, acuminate, 5–10 mm long, sparsely hairy to sub-glabrous,

longer than ovary. Ovary narrowly-obovoid, sparsely pubescent, 4–7.5 mm long. Sepals

white to light greenish, sub-similar, narrowly-ovate, obtuse, 4–5 mm long, joined on 1 / 2 –

2 /3 of their length, sparsely hairy to sub-glabrous. Petals white, narrowly-obovate,

oblique-falcate, obtuse, narrowing to the base, same length as sepals. Lip 6–7 mm

long, white, with 2 greenish spots at the base of epichile. Hypochile concave, boat

shaped, at the base with 2(3) slender finger-like recurved glands along lateral veins.

Epichile 2-lobed; lobes spreading, flabellate, 1.5–3.5 mm long and wide, with 4–7

triangular to narrowly-triangular teeth. Column 2–2.5 mm tall, with sub-equal narrowlytriangular,

acute stelidia and subulate rostellar arms. Fruit obovoid capsule to 9 mm

long. Fig. 30, j, k; 32, g.

Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.

300–1550 m. Fl. December – April. Not rare (LR).

Distribution. Vietnam (Bac Kan, Ha Giang, Lang Son, Quang Binh, Son

La), Myanmar, Hong Kong, Taiwan.

Studied specimens. Bac Kan, Cho Don, HLF 744 (HN, LE); Ha Giang, Dong Van,

HAL 8598 (HN, LE); Ha Giang, Quang Ba, DKH 4894 (HN, LE, MO), DKH 5098 (HN, LE, MO);

Ha Giang, Vi Xuyen, DKH 6309 (HN, LE, MO), DKH 6511 (HN, LE, MO); Lang Son, Bac Son,

HAL 6708 (HN, LE); Quang Binh, Bo Trach, HAL 6056 (HN, LE), HAL 6114a (HN, LE), HAL

6196 (HN, LE), HAL 6352 (HN, LE); Quang Binh, Minh Hoa, HAL 5869 (HN, LE); Son La, Moc

Chau, DKH 7509 (HN, LE, MO); Son La, Yen Chau, DKH 7111 (HN, LE, MO).

Note. Typical element of herbaceous lithophytic vegetation on mossy rocky

solid limestone.

8. C. cochinchinensis Blume, 1859, Coll. Orch. Arch. Ind. Jap.: 55; Seidenf.

1992, Opera Bot. 114: 35; Aver. 1994, Identif. Guide Vietnam. Orch.: 41; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 20. – C. flabellata auct. non Wight:

Gagnep. 1933, Fl. Gen. Indochine 6, 4: 547.

Described from southern Vietnam (“Tourane, Vietnam”). Type (“Gaudichaud

62”) – P.

Terrestrial and lithophytic herb. Stem 8–25 cm tall, with 1–4 leaves at the

base. Leaves dark green with light green median stripe, triangular-ovate to cordate,

acute, 1–3.5 cm long. Scape sparsely hairy, to 22 cm long. Rachis to 1.5 cm long,

glabrous or sparsely hairy, with 1–10 flowers. Floral bracts triangular-ovate, acuminate,

5–10 mm long, glabrous, shorter than ovary. Ovary narrowly-obovoid, glabrous, 5–

8mm long. Sepals dull greenish to pink, sub-similar, narrowly-ovate to ovate, obtuse,

4.5–6 mm long, joined on 1 / 2 – 2 / 3 of their length, glabrous. Petals white, narrowlyobovate,

oblique-falcate, obtuse, narrowing to the base, same length as sepals. Lip 8–

12 mm long, white, with triangular green papillose area at the base of epichile. Hypochile

concave, boat shaped, at the base with 3–5 slender finger-like recurved glands along


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111

lateral veins. Epichile 2-lobed; lobes spreading, flabellate, 4–6 mm long and wide,

deeply laciniate into 6–10 narrowly-triangular finger-like segments. Column 2.5–3 mm

tall, with sub-equal narrowly-triangular, acute stelidia and subulate rostellar arms. Fig.

30, l; 32, h.

Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.

700–900 m. Fl. January – February. Occasional (VU).

Distribution. Vietnam (Quang Binh, Quang Nam – Da Nang), Taiwan.

Studied specimens. Quang Binh, Bo Trach, HAL 6197 (HN); d-EXSICCATES OF

VIETNAMESE FLORA 0006/HAL6197.

Note. Typical element of herbaceous lithophytic limestone vegetation in central

Vietnam.

9. C. yunnanensis Rolfe, 1896, Kew Bull. 1896: 201; Seidenf. 1992, Opera

Bot. 114: 35; Aver. 1994, Identif. Guide Vietnam. Orch.: 41; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 20.

Described from S China (“Yunnan: ... at Mengtse”). Type (“Hancock, 25”)– K.

Terrestrial and lithophytic herb. Stem 8–30 cm tall, with 1–5 leaves at the

base. Leaves triangular-ovate to cordate, acute, 1–4 cm long. Scape tomentose, to

25cm long. Rachis to 1.5 cm long, tomentose, with 1–8 flowers. Floral bracts triangularovate,

acuminate, 5–11 mm long, tomentose, longer than ovary. Ovary narrowlyobovoid,

pubescent, 4–8 mm long. Sepals dull greenish to pink, sub-similar, narrowlyovate

to ovate, obtuse, 5–6.5 mm long, joined on 1 / 2 – 2 / 3 of their length, pubescent.

Petals white, narrowly-obovate, oblique-falcate, obtuse, narrowing to the base, same

length as sepals. Lip 8–12 mm long, white, with 2 green spots at the base of epichile.

Hypochile concave, boat shaped, at the base with 3–5 slender finger-like recurved

glands along lateral veins. Epichile 2-lobed; lobes spreading, flabellate, 4–7 mm long

and wide, with 6–10 triangular to narrowly-triangular teeth. Column 2.5–3 mm tall,

with sub-equal narrowly-triangular, acute stelidia and subulate rostellar arms. Fruit

obovoid capsule to 12 mm long. Fig. 30, m, n; 32, i.

Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.

50–1200 m. Fl. February – April. Not rare (LR).

Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Hanoi, Lang Son,

Quang Binh, Tuyen Quang), NE India, S China, Thailand.

Studied specimens. Bac Kan, Ba Be national park, HLF 510 (HN, LE); Cao Bang,

Bao Lac, CBL 1371 (HN, LE); Ha Giang, Dong Van, NTH 3526 (HN, LE); Ha Giang, Quang Ba,

DKH 4893 (HN, LE, MO), DKH 5097 (HN, LE, MO); Hai Phong, Cat Ba, LX-VN 3600 (HN, LE),

LX-VN 3426/16 (HN, LE); Lang Son, Bac Son, HAL 6709 (HN, LE); Quang Binh, Quang Ninh,

HAL 6307 (HN, LE); Quang Ninh, Ha Long Bay, NTH 5468 (HN, LE); Tonkin, Notre Dame, Tuvu,

Balansa 2003 (P); Tuyen Quang, Na Hang, NTH 2060 (HN, LE).

Note. Vicarious species replacing C. cochinchinensis in limestone areas of

northern Vietnam where it is typical element of herbaceous lithophytic vegetation on

rocky solid limestone.


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

10. C. thanmoiensis (Gagnep.) Ormerod, 2002, Lindleyana, 17, 4: 194; Aver.

et Averyanova, 2003, Updated Checklist Orch. Viet.: 20, 79. – Zeuxine thanmoiensis

Gagnep. 1931, Bull. Mus. Hist. Nat. (Paris) 2 ser. 3, 3: 327; Seidenf. 1992, Opera Bot.

114: 38; Aver. 1994, Identif. Guide Vietnam. Orch.: 38.

Described from northern Vietnam (“Tonkin: Than-moi”). Type (“19 Jan. 1886,

Balansa 313”) – P (holotype), K (isotype).

Stem 6–8 cm tall, with 3–5 leaves at the base. Leaves triangular-cordate,

acute, 1.2–1.7 cm long, petiolate. Scape pubescent, 3–4 cm long with 3–4 flowers.

Floral bracts triangular-ovate, acuminate, to 6 mm long, pubescent, shorter than ovary.

Ovary narrowly-obovoid, pubescent, to 8 mm long. Sepals sub-similar, narrowly-ovate,

obtuse, 4–5 mm long, joined on 2 / 3 of their length, hairy. Petals, narrowly-obovate,

oblique, same length as sepals. Lip 5–5.5 mm long. Hypochile concave, at the base

with 1–2 slender finger-like glands along lateral veins. Epichile entire, transverselyrhomboid,

about 1 mm long, 2.5 mm wide. Column short, with long falcate rostellum

arms and short subulate stelidia. Fig. 30, o-q.

Ecology. Not known. Very rare (DD).

Distribution. Vietnam (Lang Son).

Note. The species is known up to now by only type collection.

11. C. bipunctata Aver. 1996, Bot. Journ. (St. Petersburg), 81, 10: 78, fig. 4;

Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 20.

Described from northern Vietnam (“N Vietnam, Prov. Ha Giang, Distr. Quan

Ba, around 4 km to SW from Can Ty town.”). Type (“Averyanov et al. HG 72, 27

October 1995”) – LE.

Lithophytic herb 10–25 cm tall with 2–4 basal, shortly petiolate leaves. Leaf

blade dull silvery-green with unclear green vein network, narrowly-ovate to ovate, to

4 cm long. Scape and rachis pubescent, rachis 2–3 cm long, normally with 3–8 lax,

hardly opening flowers. Floral bracts triangular-ovate, acuminate, 4–8 mm long, not

longer than ovary, dense tomentose. Ovary 5–8 mm long, densely pubescent. Sepals

outside sparsely pubescent, light green, narrowly-ovate, 5–7 mm long, joined on 2 / 3 of

their length into a tube. Petals white or light greenish, free, lanceolate, 5–6.5 mm long,

1.5 mm wide. Lip white or light greenish, 6–7.5 mm long. Hypochile concave, basally

with numerous small glands on each side along lateral veins. Claw with 2 brightly

green spot on bended part. Epichile entire, emarginate, obcordate, or divided into 2

semi-circular lobes with entire margins. Column 1 mm tall. Anther red. Fruit obovoid,

8–10 mm long, 5–6 mm wide. Fig. 33, a-c.

Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.

500–1000 m. Fl. March – April. Rare (VU).

Distribution. Vietnam (Ha Giang).

Studied specimens. Ha Giang, Quang Ba, DKH 4895/1 (HN, LE, MO).

12. C. latipetala Aver. et Averyanova, 2006, Komarovia, 4: 5, fig. 3.

Described from northern Vietnam (“Bac Kan Province, Na Ri Distr., Na Duong


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113

Fig. 30. Cheirostylis marmorifolia: a – flowering plant and leaf, b – flower, c – sepals and

petals, d – flattened lip, e – column, frontal and dorsal view; C. filipetala: f – flowering plant

and leaf, g – flower, h – flattened tepals and lip, i – fruit and floral bract; C. chinensis: j –

flattened tepals and lip, k – column, ovary and floral bract; C. cochinchinensis: l – flattened lip;

C. yunnanensis: m – flattened tepals and lip, n – column, frontal view; C. thanmoiensis: o –

flowering plant, p – flower, q – column and lip, side view.


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

village”). Type (“25 November 2002, D.K. Harder, N.T. Hiep, L. Averyanov, DKH

7659c”) – HN (holotype), LE (isotype).

Terrestrial and lithophytic herb. Stem 5–20 cm tall, with 1–5 leaves at the

base. Leaves velvety dark green to nearly black, petiolate, triangular-ovate to cordate,

acute, to 2 cm long. Scape tomentose, to 22 cm long, with 1–4 flowers. Floral bracts

triangular-ovate, acuminate, 5–8 mm long, much shorter than ovary. Ovary narrowlyellipsoid,

sparsely pubescent, 5–6 mm long. Sepals white, glabrous, concave, obtuse, 4

mm long, dorsal sepal broadly ovate, lateral sepals narrowly-ovate, all joined on 2 / 3 of

their length. Petals white, much broadening in apical part, broadly obtuse to slightly

emarginate, 4 mm long, 1.5 mm wide. Lip 6–7 mm long, white, with 2 greenish spots

at the base of epichile. Hypochile concave, at the base with 2–3 slender finger-like

recurved glands along lateral veins. Epichile 2-lobed, 3 mm long, 6–7 mm wide; lobes

spreading, each with 3–4(5) roundish to broadly triangular blunt teeth. Column 2.5 mm

tall, with sub-equal narrow, acute stelidia and rostellar arms. Anther narrowly-ovate,

subacute, 1.5 mm long. Fig. 33, d-f; 36, a.

Ecology. Wet broad-leaved and coniferous mossy forests on rocky limestone.

400–1300 m. Fl. November – December, March. Occasional (LR).

Distribution. Vietnam (Bac Kan, Ha Giang, Son La).

Studied specimens. Ha Giang, Meo Vac, HAL 8478 (HN, LE), HAL 8526 (HN, LE);

Ha Giang, Quan Ba, HAL 8381 (HN, LE); Son La, Muong Lum, DKH 7111 (HN, LE, MO).

13. C. spathulata J.J. Smith, 1927, Bull. Jard. Bot. Buitenzorg, 3 ser. 9: 32;

Seidenf. 1992, Opera Bot. 114: 35; Aver. 1994, Identif. Guide Vietnam. Orch.: 40;

Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 20. – C. grandiflora

auct. non Blume: Gagnep. 1933, Fl. Gen. Indochine, 6, 4: 548.

Described from Java (“Java, Soerabajа: Goenoeng Kembangan bei Gresik”).

Type (“Dorgelo 2072”) – BO, L?

Stem 15–20 cm tall, with 4–6 leaves at the base. Leaves pale gray-green, with

dark gray nerves, petiolate, narrowly-ovate to ovate, acute, 1.5–2.5 cm long. Scape

12–16 cm long, tomentose, with 2–5 flowers. Sepals sub-similar, narrowly-ovate, 7–

10 mm long, joined on 1 / 2 – 2 / 3 of their length, pubescent. Petals oblique-elliptic, narrowing

to the apex an base, the same length as sepals. Lip 1.5–1.7 cm long. Hypochile concave,

boat-shaped, at the base with 4–6 small terete glands along lateral veins. Epichile 2lobed,

lobes divergent, triangular-flabellate, truncate, apex wavy, with low irregular

rounded teeth. Fig. 33, g; 36, b.

Ecology. Limestone at low elevations. Fl.? Occurrence in Vietnam needs

confirmation (EX?).

Distribution. Vietnam (probably extinct). Thailand, Java (extinct), Kalimantan.

Studied specimens. Cochinchina, Mt. Chao-lo, Pierre 145 (P).

Note. Insufficiently known species, occurrence of which in Vietnam needs

confirmation.


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Fig. 31. Corymborkis veratrifolia: a (LX-VN 3779); Tropidia angulosa: b (Averyanov 512

1988 and HAL 5261); T. curculigoides: c (DKH s.n., 1999 Ha Tinh); Anoectochilus roxburghii:

d (HAL 9735), e (HAL 3428, HAL 2303, HAL 2002); A. papillosus: f (HAL 8978, type);

Anoectochilus calcareus: g (DKH s.n., 2002, HAL 5866, HAL 1696, HAL 2994); A. albolineatus:

h (VH 4132); A. annamensis: i (HAL 8177, type).


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 32. Anoectochilus annamensis: a (HAL 6897); A. lylei: b (P 7906a); Cheirostylis serpens:

c (HAL 6092, type); C. foliosa: d (HAL 9415, type); C. takeoi: e (HAL 6708); C. marmorifolia:

f (Averyanov et al., CBL s.n., 1998, type and HLF 508); C. chinensis: g (HAL 6196 and HAL

8598); C. cochinchinensis: h (HAL 6197); C. yunnanensis: i (HAL 6389).


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117

Erythrodes Blume, 1825, Bijdr.: 410. – Physurus L.C. Rich ex Lindl. 1840,

Gen. Sp. Orch. Pl.: 501.

Type: E. latifolia Blume

Terrestrial sympodial herbs with creeping succulent rhizome rooting at nodes,

ascending erect leafy floriferous stem and terminal many flowered raceme. Leaves

few, spirally arranged at the base of stem, petiolate, sheathing at the base, glabrous,

convolute, asymmetric, uniformly green, acute. Scape hairy, leafless, with 2–4 sterile

bracts. Flowers small, resupinate. Sepals free, pubescent, lateral spreading; petals

free or connivent to median sepal forming hood. Lip lying alongside the column,

concave, saccate or spurred, not hidden by the sepals; spur or sac with 1 small flat

protuberance or flat papillose wart on each side (usually breaking off in dry herbarium

material). Epichile in form of small reflexed lamina. Column narrow, erect, apically

setaceous, acuminate, with 1 hollow stigma and 2 sectile pollinia having caudicle and

viscidium.

20(2) species. Tropical and subtropical Southeast Asia to islands of SW Pacific.

Key to species

1. Lip narrowly-oblong, boat-shaped, longer than 5 mm, about 2.5 mm wide, glabrous, at

the base with bifid spur longer than 2.5 mm; epichile sub circular, as wide as lip;

sepals 5.5–7 mm long ............................................................................................. 1. E. hirsuta

– Lip ovate, concave, 4–5 mm long, 4 mm wide, with dark papillose areas inside, at the

base saccate, sac less than 2.5 mm long; epichile broadly-obovate to triangular, as

broad as 1 / 2 of the lip; sepals 4–5.5 mm long ................................................... 2. E. blumei

1. E. hirsuta (Griff.) Ormerod, 1997, in Seidenf. Contr. Orchid Fl. Thailand,

13: 12. – Goodyera hirsuta Griff. 1851, Not. Pl. Asiat. 3: 393. – Physurus herpysmoides

King et Pantl. 1896, Journ. As. Soc. Bengal, 65: 124. – E. herpysmoides

(King et Pantl.) Schleichter, 1911, Bot. Jahrb. Syst. 45: 392. – E. seshagiriana

A.N. Rao, 1997, Indian Forester, 123: 643. – P. blumei auct. non Lindl. 1840: Gagnep.

1933, in Lecomte, Fl. Gen. Indo-Chine, 6: 567, p. p. – E. blumei auct. non (Lindl.)

Schleichter, 1905: Seidenf. 1992, Opera Bot. 114: 28; Aver. 1994, Ident. Guide Vietnam.

Orch.: 33; Aver. et Averyanova, 2003, Updated Checkl. Orch. Vietnam: 35, p. p.

Described from NE India (“Assam, on Burmese frontier, towards Nempean,

1837”). Type (“Griffith s.n.”) lost.

Stem up to 50 cm tall with 3–5 petiolate leaves. Leaf blade oblique-ovate, to

10 cm long, acute. Inflorescence glandular-pubescent, rachis to 10 cm long, with numerous

flowers. Floral bracts lanceolate, acuminate, equal or slightly longer than ovary,

8–15 mm long. Pedicel and ovary glandular-pubescent, 10–15 mm long. Sepals subsimilar,

sparsely glandular pubescent outside, narrowly-ovate, acute, spreading, 5.5–

7mm long, reddish-brown. Petals whitish, linear-spathulate, dilated at the apex, 5–

6.5 mm long, connivent to dorsal sepal forming hood. Lip red-brown, spurred, 5–6.5mm

long, oblong, boat-shaped; epichile white, small, transversely oblong or almost circular,

entire, deflexed; side-lobes small, erect. Spur short and broad, 2.5–3.5 mm long, bifid


118

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

at the apex, inside with 1 low insignificant callosity on each side. Column 4–5 mm

long. Fruit elliptic beaked capsule, 12–15 mm long. Fig. 33, h; 36, c.

Ecology. Wet closed forests, often with bamboo, on any soils. 300–500 m. Fl.

March – May. Not rare (VU).

Distribution: Vietnam (Ha Tinh, Ninh Binh, Quang Tri). Bhutan, NE India,

Myanmar, Thailand, S China.

Studied specimens. Ha Tinh, Huong Son Distr., Son Hong Municipality, HAL

5215 (HN, LE); Ninh Binh, Cuc Phuong national park, M.V. Xinh et al. MVX 173, CP 1076 (HN,

LE); LX-VN 1766, 1767 (HN, LE); Quang Tri, Da Krong, HLF 5776 (HN, LE).

2. E. blumei (Lindl.) Schleichter, 1905, in Schum. et Lauterb. Fl. Schutzgeb.

Südsee, Nachtr.: 87; Seidenf. 1992, Orch. Indochina: 28; Aver. 1994, Identif. Guide

Vietnam. Orch.: 33; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 35. –

Physurus blumei Lindl. 1840, Gen. Sp. Orch. Pl.: 504. – P. chinensis Rolfe, 1896,

Bull. Misc. Inf. Kew, 1896: 200. – E. chinensis (Rolfe) Schleichter, 1914, Die

Orchideen: 117.

Described from NE India (“India orientali; provincia Sylhet...”). Lectotype

(“Wallich 7397”) – K.

Stem up to 60 cm tall with 3–4 petiolate leaves. Leaf blade oblique-ovate, to

12 cm long, acute. Inflorescence glandular-pubescent, rachis to 15 cm long, with

numerous flowers. Floral bracts lanceolate, acuminate, 4–10 mm long. Pedicel and

ovary sparsely pubescent with glandular hairs to subglabrous, to 14 mm long. Sepals

sub-similar, subglabrous or sparsely glandular pubescent outside, narrowly-ovate, obtuse,

spreading, 4–5.5 mm long, brownish-green with central brown-purple spot. Petals

whitish with brown-purple median vein, lanceolate-spathulate, dilated at the apex, 3.5–

5 mm long, connivent to dorsal sepal forming hood. Lip pinkish-brown, saccate, 4–

5mm long, obovate, concave; epichile white, broadly-obovate to triangular, entire,

deflexed; side-lobes, erect, hemicircular. Sac short, cup-shaped, 1.5–2.5 mm long,

inside with 1 low papillose circular gland. Column 3–4 mm long. Fig. 33, i; 36, d.

Ecology. Wet closed forests, often with bamboo, on any soils. 400–1400 m.

Fl. March – May. Not rare (VU).

Distribution: Vietnam (Bac Kan, Ha Giang, Ha Tinh, Hoa Binh, Lao Cai,

Thua Thien Hue, Vinh Phuc, Yen Bai), NE India, Myanmar, Thailand, S China, Taiwan,

Malaya, Sumatra, Java.

Studied specimens. Ba Be, Bac Kan, Eberhardt s.n.; Ha Giang, Bat Dai Son mt.,

DKH 5073 (HN, LE, MO); Ha Tinh, Huong Son, HAL 5215 (HN, LE); Hoa Binh, Da Bac, HAL 557

(HN, LE); Lao Cai, Van Ban, HAL 2266 (HN, LE); Thua Thien Hue, Bach Ma national park, HLF

904 (HN, LE); Vinh Phu, Tam Dao, a.1987, Averyanov, LX-VN s.n. (LE); Yen Bai, Regnier s.n.

Goodyera R. Br. 1813, Hort. Kew. 2 ed. 5: 197.

Type: G. repens (L.) R. Br. (Satyrium repens L.).

Terrestrial, lithophytic and occasionally epiphytic sympodial creeping herbs

with few rosette leaves and ascending floriferous leafless stem. Leaves petiolate,

sheathing at base, acute. Scape erect, with few sterile bracts and 1 to many resupinate


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119

flowers in more or less dense, often secund spike. Sepals all parallel to the floral axis,

or 2 lateral sepals spreading. Petals oblique, usually connivent and forming hood with

median sepal, sometimes joined near their apex. Lip divided into hypochile and epichile.

Hypochile saccate, with bristly hairs or lamellar protuberance inside. Epichile simple,

entire, oblong-ligulate, usually reflexed. Column short, without appendages at base.

Rostellum long, deeply cleft into long arms. Pollinia 2, sectile, with caudicle and elongate

viscidium. Stigma entire, hemispheric, on the front of the column.

80–100(9) species. Europe, Mainland Asia, Indonesia, N Australia, islands of

W Pacific, N Australia, North and Central America.

Key to species

1. Rhizome very short, floriferous stem erect; leaves more or less erect, broadly-lanceolate

to narrowly ovate; scape, rachis and flowers glabrous; inflorescence dense many

flowered spike; sepals 2–3 mm long ................................................................... 1. G. procera

– Rhizome more or less long, creeping, floriferous stem ascending; leaves spreading,

narrowly-ovate to broadly-ovate; scape, rachis and flowers mоre or less hairy;

inflorescence lax or sub-dense, few- to many-flowered spike; sepals longer than 3 mm ...... 2

2. Large plants, with inflorescence 0.5–1 m tall; spike with more than 30 flowers; leaves

10–20 cm long, creeping stem 5–10 mm thick .................................................... 2. G. fumata

– Plants smaller, with inflorescence less than 0.5 m tall; spike with less than 30 flowers;

leaves less than 10 cm long; creeping stem less than 5 mm thick .................................. 3

3. Leaves distinctly reticulated with white, yellow or pink veins ...................................... 4

– Leaves uniform green, sometime with 1 median white stripe or mottled with white

irregular spots ...................................................................................................................... 5

4. Inflorescence many-flowered spike 3–7 cm long on scape longer than 5 cm; sepals 3–

3.5 mm long, densely tomentose ......................................................................... 3. G. hispida

– Inflorescence subsessile with 1–3 flowers on scape less than 1 cm long; sepals 18–

22mm long, glabrous or sparsely hairy ............................................................... 4. G. biflora

5. Inflorescence with 1–3 flowers 2–2.5 cm across, sepals glabrous, spreading, 12–

14mm long ........................................................................................................ 5. G. viridiflora

– Inflorescence normally with 4–25 flowers, flowers less than 1 cm across, sepals

pubescent or sparsely hairy, not spreading, less than 13 mm long ............................... 6

6. Floral bracts exceeding half of flowers, longer than 1.5 cm; sepals usually longer than

10 mm; leaves 2.5–10 cm long ............................................................................... 6. G. foliosa

– Floral bracts not exceeding half of flowers, shorter than 1.5 cm; sepals usually less

than 10 mm long; leaves 1.5–4 cm long ............................................................................. 7

7. Sepals 8–9 mm long, pubescent; leaves mottled with white irregular spots, rare pure

green ................................................................................................... 7. G. schlechtendaliana

– Sepals less than 8 mm long, glabrous or sparsely hairy; leaves uniform dark green to

dark olive-green, sometimes with white median stripe .................................................... 8

8. Leaves dark uniform green; inflorescence dense secund spike with 10–25 flowers;

hypochile glabrous, with 2 lamellate parallel protuberance near the base .... 8. G. arisanensis

– Leaves dark green to dark olive-green, with white median stripe; inflorescence lax

spike, with few spirally arranged flowers; hypochile pilose with numerous bristly

hairs ............................................................................................................ 9. G. rhombodoides


120

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

1. G. procera (Ker-Gawl.) Hook. 1823, Exot. Fl. 1, 3, tab. 39; Seidenf. 1992,

Opera Bot. 114: 38; Aver. 1994, Identif. Guide Vietnam. Orch.: 36; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 37.– Neottia procera Ker-Gawl. 1822, Bot.

Reg. 8, tab. 639.

Described from Nepal (“Nepal”). Type – icon (“Wallich 619”) – K.

Erect terrestrial and lithophytic herb up to 60 cm tall, with 4–8 petiolate erect

leaves in basal third. Leaves broadly-lanceolate to narrowly-ovate, 5–15 cm long, 1—

4 cm wide, pure light green. Scape glabrous, with 2–4 sterile narrow, acuminate bracts

and dense, many-flowered spike 5–20 cm long; scape finely papillose. Floral bracts

glabrous, ovate-lanceolate, acuminate, 5–8 mm long, ciliate along margin. Ovary

glabrous, 3–5 mm long. Flowers hardly opening, sub-globose, 3–4 mm across. Sepals

similar, concave, ovate, 2–3 mm long, glabrous, light greenish, brownish or olive-green.

Petals white, as long as sepals, oblique-spathulate. Lip white, sub-globose, saccate,

1.5–2 mm long, with few slender glands within; apex triangular, blunt recurved, with 2

fleshy tubercles at the base. Column 1–1.5 mm high, with acute rostellar arms, and

ovate, acute anther. Fruit ovoid capsule, 4–5 mm long. Fig. 33, j, k; 36, e.

Ecology. Wet open mossy places among secondary grasslands and open shrubs,

commonly along rocky stream valleys and on spring seepages, on any kinds of soils.

100–1600 m. Fl. February – May. Common (LR).

Distribution. Vietnam (probably all provinces), Sri Lanka, Nepal, Bhutan,

India, Japan, China, Taiwan, Hainan, Myanmar, Thailand, Laos, Cambodia, Malaysia,

Java, Sumatra, Philippines.

Studied specimens. Bac Kan, Ngan Son, CBL 1259 (HN, LE); Binh Tri Thien, Bogiang,

Dak Lac, Krong Bong, VH 6461 (HN, LE); Gia Lai – Kontum, Dakgley, Bien 565 (HN,

LE), Phoong 426 (HN, LE); Gia Lai – Kontum, Kbang, LX-VN 2085 (HN, LE); Gia Lai – Kontum,

Kon Ha Nung, LX-VN 1991 (HN, LE); Ha Giang, Bac Me, HAL 6608 (HN, LE); Ha Giang,

Quang Ba, DKH 4834 (HN, LE), Bien 14248 (LE); Hoang Lien Son, Sapa, 19.5.61, sine coll.,

2312, 2318 (LE); Kon Tum, Chu Mom Ray, DKH 4553 (HN, LE); Kon Tum, Ngoc Linh mt, VH

297 (HN, LE), VH 1190 (HN, LE); Lai Chau, Muong Te, HAL 10120 (HN, LE); Lai Chau, Phong

Tho, HAL 10713 (HN, LE); Ninh Thuan, Nui Chua mt., HLF 4179 (HN, LE); Quang Tri, Da

Krong, HLF 5756 (HN, LE), HLF 6163 (HN, LE); Quang Tri, Huong Hoa, HLF 5901 (HN, LE),

d-EXSICCATES OF VIETNAMESE FLORA 0080/HLF5901; Thua Thien – Hue, A Luoi, HAL

7266 (HN, LE), HAL 7502 (HN, LE); Thua Thien – Hue, Bach Ma national park, HLF 1732 (HN,

LE); Thua Thien – Hue, Huong Thuy, HAL 8221 (HN, LE); Thua Thien – Hue, Nam Dong, HAL

6924 (HN, LE), HAL 10867 (HN, LE); Thuan Hai, Ka-rom.

2. G. fumata Thwaites, 1861, Enum. Pl. Zeylan.: 314; Seidenf. 1992, Opera

Bot. 114: 28; Aver. 1994, Identif. Guide Vietnam. Orch.: 36; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 37. – Allochilus eberhardtii Gagnep. 1932,

Bull. Mus. Hist. Nat. 2 ser. 4: 591.

Described from Sri Lanka (“Ceylon... Central Province”). Type (“Thwaites

CP 3668”) – K (holotype), PDA (isotype).

Terrestrial and lithophytic herb with ascending fleshy leafy stem 6–10 mm


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121

thick and terminal inflorescence 50–100 cm tall. Leaves 3–7; petiole 2–6 cm long;

leaf blade oblique-elliptic, acute, 10–20 cm long, 4–8 cm wide, pure green. Scape

leafless, pubescent, with few narrow, acuminate sterile bracts, bearing dense, manyflowered

spike 15–45 cm long. Floral bracts cuneate, 0.5–1 cm long. Flowers widely

opening, light brownish to olive-greenish, lip white. Ovary densely glandular pubescent,

1–1.2 cm long. Sepals sub-similar, narrowly ovate, obtuse, pubescent, 6–8 mm long;

lateral sepals reflexed. Petals oblique lanceolate-spathulate, dilated to the apex, as

long as sepals. Lip concave, cup-shaped, 5–6 mm long, 4–5 mm wide, hairy inside,

with attenuate narrowly-oblong, obtuse, much reflexed apex. Column stalked dilated

above with emarginate rostellum, 4–5 mm long. Fruits obovoid capsule, 1–1.2 cm

long, 4–5 mm wide. Fig. 33, l; 36, f.

Ecology. Wet closed forests on any kind of soils, commonly along humid

rocky stream canyons. 400–2000 m. Fl. January–April. Occasional (LR).

Distribution. Vietnam (Bac Kan, Dak Nong, Kon Tum, Lam Dong, Ninh

Binh), Sri Lanka, India, Myanmar, China, Ryukyu Islands, Taiwan, Thailand, Malaysia,

Philippines, Java.

Studied specimens. Bac Kan, Ba Be, HLF 555 (HN, LE); Bac Kan, Cho Don, HAL

4696 (HN, LE); Bac Thai, Chu-bo; Siam-Khong, Dak Nong, Dak Glong, HLF 5610 (HN, LE); Ha

Nam Ninh, Cuc Phuong, 1989, Bao 631 (HN, LE), 20.03.1971, Trong 155 (HN, LE), N.M. Cuong,

H.L. Quyen, NMC 289 (HN), D.D. Soejarto et al. DDS 11264 (HN); sine date, MAJ 514 (HN);

Ha Nam Ninh, Cuc Phuong, May bac, LX-VN 1811 (HN, LE); Kon Tum, Dak Gley, VH 2344

(HN, LE); Kontum, Ngoc Linh mt., VH 833 (HN, LE); Lam Dong, Lac Duong, VH 3329 (HN, LE).

Note. Widespread and common species growing in shady wet woods at different

elevations, commonly along stream and wet valleys. Rarely observed in flowers.

Typical herbaceous element of humid forest understory.

3. G. hispida Lindl. 1857, Journ. Linn. Soc. London (Bot.), 1: 183; Seidenf.

1992, Opera Bot. 114: 29; Aver. 1994, Identif. Guide Vietnam. Orch.: 36; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 37.

Described from NE India (“... Khasia Hills”). Type (“Hooker & Thomson

2110”) – K.

Terrestrial and lithophytic herb 6–20 cm tall. Leaves 2–5, petiolate; petiole

0.6–3 cm long; leaf blade ovate to lanceolate, acuminate, 4–14 cm long, 1.2–2.7 cm

wide, velvety dark brown-green, reticulated with silvery, yellow or pinkish veins, often

with light broad median stripe. Scape pubescent, with 2–4 sterile narrow, acuminate

bracts and dense, many-flowered spike. Rachis hairy, 3–7 cm long. Floral bracts

glabrous to sparsely hairy, lanceolate, acuminate, 7–11 mm long. Ovary densely hairy,

4–6 mm long. Flowers hardly opening. Sepals ovate, concave, 3–4 mm long, 1–1.5 mm

wide, hairy, olive-green, with white tips. Petals white, of the same length, obliqueoblong,

to lanceolate, with wavy obtuse apex. Lip white, concave, cup-shaped, 2–

3mm long, 2 mm wide, coarsely pubescent within, with attenuate recurved apex.

Column 1 mm high, with linear rostellar arms. Fruit broadly elliptic to almost round

capsule, 5–8 mm long. Fig. 33, m-o; 36, g.


122

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Ecology. Closed forests on any kind of soils. 150–1500 m. Fl. June – August.

Not rare (VU).

Distribution. Vietnam (probably all provinces), India, Thailand, Peninsular

Malaysia.

Studied specimens. Bac Kan, Cho Don, HAL 4700 (HN, LE), HLF 760 (HN, LE);

Bac Kan, Na Ri, HAL 4934 (HN, LE); Cao Bang, Bao Lac, CBL 310 (HN, LE); Cao Bang,

Nguyen Binh, CBL 1304b (HN, LE); Dak Lac, Chu Yang Sinh mt., VH 6102 (HN, LE); Ha Giang,

Bac Me, HAL 6574 (HN, LE), HAL 6421 (HN); Ha Giang, Quang Ba, DKH 4788 (HN, LE), DKH

4823 (HN, LE); Ha Giang, Vi Xuyen, DKH 6346 (HN, LE, MO); Ha Nam Ninh, Cuc Phuong,

Luan Van Bao 707 (HN, LE), LX-VN 1808 (HN, LE), 23 July 1965, Bich et al. 948, CPNP 5149

(HN), 25 March 1971, Hien B-50, CPNP 5123 (HN), 17 July 1971, Hop, Khoi 565, CPNP 5122

(HN), 12 July 2002, NMC 1680 (HN), sine date, Aung 784 (HN), sine date, Bich, Nhu, Liem 948

(HN), sine date, Dung 784, CPNP 5124 (HN), sine date, Hien B-51(HN), sine date, N.T. Hiep,

NTH 3892 (HN), sine date, Hop, Khoi 565 (HN); Hai Phong, Cat Ba, LX-VN 3446 (HN, LE);

Khanh Hoa, Hon Giao, 1997, VH s.n. (LE – photo); Ninh Binh, Cuc Phuong, HAL 1659 (LE –

photo); Ninh Thuan, Nui Chua national park, HLF 4286 (HN, LE); Quang Binh, Phong Nha-Ke

Bang national park, HAL 6189 (HN, LE – photo); Son La, Moc Chau, HAL 9385 (HN); Son La,

Yen Chau, HAL 9463 (HN); Thanh Hoa, Ba Thuoc, HAL 337 (HN, LE), HAL 977 (HN), HAL

3010 (HN, LE), HAL 3182 (HN), HAL 3312 (HN, LE), HAL 3371 (HN), Thanh Hoa, Co Lung,

2003, HAL 3312 (LE – photo); Thua Thien – Hue, Bach Ma national park, HLF 1624 (HN, LE),

Lai Chau, Sin Ho, HAL 9881 (HN, LE).

Note. Typical shade-loving element of forest understory. It is easily distinguished

by its deep velvety brown-green leaves reticulated with silvery, yellow or pink veins.

The species is widely collected for traditional oriental medicine and as ornamental plant

known under common name – “jewel orchids”. Needs special attention for protection.

4. G. biflora (Lindl.) Hook. f. 1890, Fl. Brit. India 6: 114. – Georchis biflora

Lindl. 1840, Gen. Sp. Orch. Pl.: 496.

Described from Nepal? Type (“Wall. Cat. 7379”) – K.

Terrestrial and lithophytic herb 5–8 cm tall. Leaves 4–6, shortly petiolate; leaf

blade ovate to broadly-ovate, acute, 3.5–5 cm long, velvety dark green, reticulated

with silvery veins. Inflorescence sub-sessile with hairy scape less than 1 cm long and

1–3 terminal secund flowers. Floral bracts glabrous to sparsely hairy, ovate to obovate,

shortly acute, 0.9–1.4 cm long. Ovary sessile, hairy, 8–12 mm long. Flowers odorless,

not widely opening. Sepals oblanceolate, concave, 1.8–2.2 cm long, sparsely hairy,

white to light pink. Petals white, of the same length, oblique-rhomboid, with finely

plicate margin. Lip white, oblong-ligulate, 1.8–2 cm long, with concave, coarsely hairy

base and recurved acute apex plicate along margin. Column erect 2.5–3.5 mm tall;

linear rostellar arms and narrowly cuneate anther close each other, to 1 cm long. Fig.

33, p-r; 36, h.

Ecology. Primary evergreen broad-leaved wet forests on silicate rocks. 1500–

1800 m. Fl. April – May. Very rare (VU).

Distribution. Vietnam (Lao Cai), Nepal, NE India, Japan, Korea, China,

Taiwan.


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123

Fig. 33. Cheirostylis bipunctata: a – flowering plant and leaf, b – flower, c – flattened tepals

and lip; C. latipetala: d – flowering plant, e – flattened tepals, flattened lip and column, side

view, f – ovary; C. spathulata: g – flowers and base of spur; Erythrodes hirsuta: h – lip, frontal

view; E. blumei: i – flattened tepals, column and lip; Goodyera procera: j – flower and lip, k –

column, side view; G. fumata: l – flattened tepals, flattened lip and column, half-side view;

G. hispida: m – flowering plant, n – flower without lip and petals, o – lip; G. biflora: p –

flattened tepals and lip, q – column, side view, r – floral bract; G. viridiflora: s – flattened flower

without petals and lip, flattened lip and flattened petals.


124

Studied specimens. Lao Cai, Sa Pa, HAL 11135 (LE).

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

5. G. viridiflora (Blume) Blume, 1858, Coll. Orchid.: 41; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 37. – Neottia viridiflora Blume, 1825, Bijdr.: 408.

Described from Java (“Java, Salak”). Type (“Blume s.n.”) – P.

Terrestrial and lithophytic herb 12–20 cm tall with 3–4 shortly petiolate rosette

leaves. Leaf blade dull green, ovate, with cordate base, 2–9 cm long. Scape 4–15 cm

long, pubescent, with 3–4 large, lanceolate, acuminate, sterile bracts, 2–3 cm long and

1–3 pale green to light pink-brown flowers. Floral bracts triangular-ovate, acuminate,

glabrous, ciliate along margin, 1.5–2 cm long. Ovary glabrous, 8–10 mm long. Sepals

glabrous, narrowly-ovate, 12–14 mm long; lateral sepals spreading. Petals obliquerhombic,

dilated at the middle, same length as sepals, connivent to median sepal forming

hood. Lip ovate, as long as sepals, with concave hairy base and attenuate, acute apex

turned down. Column 2–3 mm long, widened at the stigma, bearing a very long rostellum

to 10 mm long, with long beaked anther behind it. Fruit ovoid capsule to 1.2–1.4 cm

long. Fig. 33, s; 36, i.

Ecology. Broad-leaved and coniferous mossy forests, mainly on rocky

limestone. 500–2000 m. Fl. March – April. Rare (VU).

Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Lao Cai, Quang Binh,

Son La), India, Nepal, Bhutan, China, Ryukyu Islands, Thailand, Malaysia, Philippines,

Indonesia to New Guinea and Samoa.

Studied specimens. Bac Kan, Cho Don, HAL 4865 (HN, LE), HLF 768 (HN, LE);

Cao Bang, Yen Lac, 25 Dec. 1998, Averyanov et al. s.n. (LE); Ha Giang, Quang Ba, DKH 5278

(HN, LE); Lao Cai, Van Ban, HAL 2509 (HN, LE); Quang Binh, Bo Trach, HAL 6061 (HN, LE);

Quang Binh, Minh Hoa, HAL 6014 (HN, LE); Son La, Moc Chau, DKH 7400 (HN, LE).

6. G. foliosa (Lindl.) Benth. ex Clarke, 1889, Journ. Linn. Soc. London (Bot.)

25: 73; Seidenf. 1992, Opera Bot. 114: 29; Aver. 1994, Identif. Guide Vietnam. Orch.:

35; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 37. – Georchis foliosa

Lindl. 1840, Gen. Sp. Orch. Pl.: 406.

Described from Myanmar (“in regno Burmano”). Type (“Griffith s.n.”) – K.

Creeping terrestrial herb 15–35 cm tall. Leaves 4–6, petiolate, oblique-elliptic,

acute, 4–10 cm long, 1.5–4 cm wide, pure green, often finely undulate along margin.

Petiole 1–4 cm long. Scape leafless, with dense, many-flowered slightly secund spike.

Rachis sparsely hairy, 1.5–10 cm long. Floral bracts ovate to lanceolate, acuminate,

glabrous, 1.5–2 cm long, as long as half of flowers or longer. Ovary densely hairy, 5–

12 mm long. Flowers odorless, not widely opening. Sepals sub-similar, olive-brown to

pinkish, ovate, concave, obtuse, 9–13 mm long, pubescent. Petals white, oblique

rhomboid-spathulate, 8–12 mm long, 2–3 mm wide, forming hood with dorsal sepal.

Lip white to light greenish or pinkish, ovate, 7–10 mm long, 4–5 mm wide, with concave,

cup-shaped, long-papillose base and attenuate, obtuse white recurved obtuse apex.

Column 3–3.5 mm tall, with straight, finger-like rostellum arms and narrowly ovate

attenuate anther 4–4.5 mm long. Fruit ellipsoid capsule, 10–14 mm long, 4–5 mm


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wide. Fig. 34, a; 37, a.

Ecology. Wet closed forests on any kinds of soil. 400–2000 m. Fl. September –

November. Not rare (LR).

Distribution. Vietnam (Bac Kan, Cao Bang, Dak Nong, Ha Giang, Kon Tum,

Lai Chau, Lam Dong, Lao Cai, Ninh Binh, Ninh Thuan, Thanh Hoa, Thua Thien –

Hue, Vinh Phu), Bhutan, Nepal, NE India, Myanmar, Japan, Korea, China, Taiwan,

Thailanad.

Studied specimens. Bac Kan, Ba Be national park, HLF 614 (HN, LE); Bac Kan,

Cho Don, HAL 4870 (HN, LE); Cao Bang, Pho Bang, Dak Nong, Dak Glong, HLF 5611 (HN,

LE); Ha Giang, Bac Me, HAL 6586 (HN, LE); Ha Giang, Meo Vac, NTH 3402 (HN, LE); Ha Nam

Ninh, Cuc Phuong, Bao 631 (LE); Kon Tum, Dak Gley, VH 1674 (HN, LE); Kon Tum, Dak Gley,

VH 2334 (HN, LE); Kon Tum, Dak Gley, VH 2406 (HN, LE); Lai Chau, Sin Ho, HAL 10027 (HN,

LE); Lam Dong, 14 km to NNW from Dalat city, VH 2522 (HN, LE); Lam Dong, Lac Duong, VH

2634 (HN, LE); Lam Dong, Lac Duong, VH 3126 (HN, LE); Lao Cai, Van Ban, HAL 2828 (HN,

LE); Ninh Binh, Cuc Phuong, 8 Dec. 2001, N.T. Hiep s.n. (HN, LE); Ninh Thuan, Ninh Son, VH

3627 (HN, LE); Thanh Hoa, Ba Thuoc, HAL 3967 (HN, LE); Thua Thien – Hue, Xuan Loc, HLF

1826 (HN, LE); Vinh Phu, Tam Dao, Hiep, 14900 (LE); Vinh Phu, Tam Dao, LX-VN 2356 (HN, LE).

Note. Widespread and common species growing in shady wet woods at different

elevations, but rarely flowers. Typical herbaceous element of forest understory.

7. G. schlechtendaliana Reichenb. f. 1849, Linnaea 22: 861; Seidenf. 1992,

Opera Bot. 114: 29; Aver. 1994, Identif. Guide Vietnam. Orch.: 35; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 37.

Described from Japan (“Japan”). Type (“Göring s.n.”) – W.

Terrestrial, lithophytic and occasionally epiphytic herb with 2–6 rosette leaves

and ascending floriferous stem 10–25 cm tall. Leaves petiolate, ovate, acute, 2–6 cm

long, light green, with white mottling or irregular spots. Scape and rachis densely

pubescent. Spike 5–12 cm long, secund, with 5–15 lax to sub-dense flowers. Floral

bracts cuneate, acuminate, 6–11 mm long, longer than ovary, hairy. Ovary ellipsoid, 5–

11 mm long, white glandular pubescent. Flowers white, sometimes with greenish or

pink tint, apices of sepals and lip sometimes with dark green spot. Sepals sub-similar,

ovate, acute, 8–9 mm long, with 1 prominent vein, densely white glandular pubescent;

lateral sepals oblique, with reflexed apex. Petals oblique-rhomboid, same length as

sepals, finely wavy along margin toward the apex. Lip 6–9 mm long, with saccate,

concave, pilose base and oblong-ligulate, attenuate, down curved blunt apex. Column

straight, 2.5–3 mm tall, with subulate rostellar arms 4–5.5 mm long and ovate anther.

Fruit ellipsoid capsule, 8–12 mm long. Fig. 34, b; 37, b, c.

Ecology. Wet mossy cloudy forests and bamboo-ericaceous thickets on mountain

tops on any kinds of soils. 1300–3000 m. Fl. September – December. Common (LR).

Distribution. Vietnam (probably all montane regions with elevations higher

than 1300 m), NE India, Nepal, Bhutan, China, Taiwan, Korea, Japan, Thailand, Sumatra.

Studied specimens. Cao Bang, Bao Lac, CBL 311 (HN, LE); Cao Bang, Nguyen

Binh, CBL 1580 (HN, LE); Ha Giang, Dong Van, CBL 1799 (HN, LE), NTH 3370 (HN, LE); Ha

Giang, Quan Ba, HAL 8391 (HN); Ha Giang, Yen Minh, HAL 8404 (HN, LE), Yen Minh, NTH


126

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

3495 (HN, LE); Hoa Binh, Mai Chau, DKH 7923 (HN, LE, MO); Kontum, Ngoc Linh mt., VH

581 (HN, LE); Lai Chau, Tam Duong, HAL 10186 (HN), HAL 10341 (HN, LE); Lam Dong, 14 km

to NNW from Dalat, VH 2576 (HN, LE); Lam Dong, Bi Doup – Nui Ba national park, HLF 5257

(HN, LE); Lam Dong, Lac Duong, Bi Doup mt., HLF 5257 (HN, LE), d-EXSICCATES OF

VIETNAMESE FLORA 0038/HLF5257; Lam Dong, Lac Duong, Bi Dup mt., VH 3304 (HN,

LE), VH 4454 (HN, LE); Lam Dong, Lac Duong, VH 2868 (HN, LE); Lam Dong, peak of Bi Dup

mt., VH 2999 (HN, LE); Lao Cai, Fan Si Pang mt., NTH 2317a (HN, LE).

8. G. arisanensis Hayata, 1916, Icon. Pl. Formos. 6: 91.

Described from Taiwan (“Mt. Arisan, ad 2500 m alt.”). Type (“U. Faurie,

Dec. 1914“) – TI?

Terrestrial herb with 2–4 rosette leaves and ascending floriferous stem 5–12

cm tall. Leaves petiolate, narrowly-ovate, dark green, acute, 2–3.5 cm long. Scape

and rachis pubescent. Spike dense flowered, secund, with 10–25 flowers. Floral bracts

cuneate, acuminate, 4–10 mm long, longer than flowers, glabrous, ciliate along margin.

Ovary ellipsoid, to 6 mm long, sparsely hairy or glabrous. Sepals narrowly-ovate, obtuse,

4–5 mm long, with 1 prominent vein, lateral sepals oblique. Petals oblanceolate, falcateoblique,

same length as sepals. Lip 4–5 mm long, saccate at the base and oblongligulate,

blunt apex. Hypochile concave, protruding outside lateral sepals, glabrous, but

with 2 lamellate parallel protuberance near base. Column 1.5–2 mm tall, anther ovate,

rostellum with 2 slender arms. Fruit ellipsoid capsule, 6–8 mm long, 3–4.5 mm wide.

Fig. 34, c, d.

Ecology. Primary broad-leaved cloudy forests and ericaceous thickets on

tops of mountains composed with silicate rocks. 2300–2500 m. Fl. October – December.

Rare (VU).

Distribution. Vietnam (Lai Chau), Taiwan.

Studied specimens. Lai Chau, Tam Duong, HAL 10468 (HN, LE).

9. G. rhombodoides Aver. 2008, Taiwania, 52, 4: 297, fig. 8.

Described from northern Vietnam (“Lai Chau Prov., Tam Duong Distr. ...”).

Type (“3 Dec. 2006, Nguyen Tien Hiep, L. Averyanov, Pham Van The, HAL 10338”) –

HN (holotype), LE (isotype).

Terrestrial herb 12–15 cm tall, with 3–5 petiolate leaves at the base. Leaves

oblique, ovate, acute, 1.5–2.5 cm long, 0.8–1.4 cm wide, velvety dark green to olivegreen,

with white median stripe. Scape 3–5 cm long, short densely hairy, with 2–3

cuneate, acuminate, sub-glabrous, sterile bracts 5–8 mm long and lax spike with 3–6

flowers. Floral bracts cuneate, acuminate, sub-glabrous or sparsely hairy, 6–10 mm

long. Ovary cylindric, 4.5–6 mm long, 0.8–1 mm wide, short densely hairy. Flowers

not widely opening. Sepals sub-similar, narrowly-ovate, sub-glabrous, 6–7 mm long,

2–2.5 mm wide, lateral sepals cucullate, reflexed at a right angle. Petals oblique,

lanceolate-rhomboid, as long as sepals, 1–1.4 mm wide, connivent with the dorsal

sepal forming forward directed hood. Lip 5.5–6.5 mm long, 2.5 mm wide; base

hemispheric, cup–like, densely hairy with long hairs; apex entire, oblong-ligulate, slightly


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concave, obtuse to blunt. Column 2.5–3 mm tall, with 2 short rostellar wings. Anther

narrowly-ovate, acuminate, 1.6–1.8 mm long. Fig, 34, e–h.

Ecology. Primary broad-leaved evergreen forests on granite and shale. 1900–

2400 m. Fl. September – November. Very rare (DD).

Distribution. Vietnam (Lai Chau).

Note. Very rare species known by the only type collection. On floral structure

it resembles Goodyera foliosa, but distinctly differs in twice smaller flowers, small

size of plant and in small, dark velvety green leaves with distinct white longitudinal

median stripe.

Herpysma Lindl. 1833, Bot. Reg. 19, tab. 1618.

Type: H. longicaulis Lindl.

Terrestrial sympodial herbs 20–35 cm tall, with creeping succulent rhizome

rooting at nodes, ascending erect leafy floriferous stem and terminal short, dense

spike. Leaves narrowly-ovate, 5–9 cm long, acute, distant, spirally arranged, petiolate,

sheathing at the base, glabrous, convolute, acuminate. Scape short, pubescent. Rachis

2–3 cm long with several dense flowers. Floral bracts narrowly-ovate, obtuse, exceeding

the ovary, pubescent. Ovary 8–10 mm long. Flowers sessile, resupinate, white with

pink tint or pink median veins on tepals and lip. Sepals sub-similar, free, narrowlyovate,

acute, concave, pubescent, 7–11 mm long. Petals rhomboid, acute, as long as

median sepal, joined in upper half and connivent to median sepal forming hood. Lip

spurred; blade rectangular, 6–9 mm long, deflexed from the middle, with rounded side

lobes and rectangular median lobe with median dent; disk with 2 short triangular keels

between side lobes and 1 longitudinal keel on mid lobe. Spur slender, cylindrical, 8–

9mm long, bifid at apex, inside with several small scattered warts. Column adnate to

basal lip margins, 5–6.5 mm tall, wingless, with bifid, rather short rostellum and large

entire stigma at front. Anther ovate. Pollinia 2, sectile, with caudicle and viscidium.

Fig. 34, i-l.

Monotypic genus.

1. H. longicaulis Lindl. 1833, Bot. Reg. 19, tab. 1618; Seidenf. 1992, Orch.

Indochina: 28; Aver. 1994, Identif. Guide Vietnam. Orch.: 33; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 40.

Described from Nepal (“Napalia”). Type (“Wallich 7389”) – K, C.

Ecology. Broad-leaved forests on silicate soils. 1500–2000 m. Fl. September –

November. Very rare (DD).

Distribution. Vietnam (Lam Dong), Nepal, Bhutan, NE India, Myanmar, S

China, Thailand, Sumatra.

Studied specimens. Annam, sine loc, Vernet, 12 (P); Dankia, Langbian, Poilane,

18666 (P).

Note. Very rare species not observed in Vietnam during last decades.

Hetaeria Blume, 1828, Fl. Jav. Orch. 1 Pref.: 7, nom. emend. id. 1858, Coll.


128

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Orch. Arch. Ind. Jap.: 102, tab. 32; Seidenf. 1978, Dansk Bot. Ark. 32, 2 : 94–104. –

Etaeria Blume, 1825, Bijdr. 6, fig. 14; id. 1825, ibid. 8: 409, orth. var. rej.

Lectotype: H. oblongifolia (Blume) Blume (Etaeria oblongifolia Blume)

Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome

rooting at nodes, ascending erect leafy floriferous stem and terminal many-flowered

raceme. Leaves few, spirally arranged, petiolate, sheathing at the base, glabrous,

convolute, asymmetric, uniformly green, acute. Scape leafless, with 2–4 sterile bracts.

Flowers small, not or hardily resupinate. Sepals free; petals free or connivent to median

sepal forming hood. Lip spurless. Hypochile concave with various papillae on lateral

sides. Epichile straight, acute or reflexed in form of small circular plate. Column short,

erect, with prominent rostellar arms and two wing-like appendages at front. Anther

narrowly triangular-ovate, acute. Pollinia 2, sectile, with stipe and viscidium. Stigma

with 2 convex, separated lobes.

30(5) species. Tropical Africa and Asia, Australia, Pacific islands to Tahiti.

Key to species

1. Epichile of the lip minute, less than 5 mm wide, heart-shaped, triangular acute or

obtuse .................................................................................................................................... 2

– Epichile of the lip large, 6–8 mm wide, with large orbicular upwards-backwards turned

white lobules ....................................................................................................... 1. H. anomala

2. Wings of column incurved or folded, not straight, not reaching apex of rostellar arms ....... 3

– Wings of column narrow, straight, widening into apical spathulate prolongation as

long as rostellar arms, or longer ........................................................................... 2. H. alta

3. Hypochile with 5–6 slender papillae placed on each side of the sac .... 3. H. oblongifolia

– Hypochile with 1–3 fat irregularly branching or flattened irregularly dented glands on

each side of the sac ............................................................................................................. 4

4. Sepals 4–5 mm long; petals narrowly-obovate, hammer-shaped, very oblique, widening

from narrow base into semicircular widening on upper edge faced to dorsal sepal ........

.................................................................................................................................... 4. H. affinis

– Sepals 5–6 mm long; petals ovate to broadly-ovate, without semicircular widening,

not oblique or slightly oblique .............................................................................. 5. H. nitida

1. H. anomala (Lindl.) Reichenb. f. 1874, Trans. Linn. Soc. Bot. 30: 142;

Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 40. – Aetheria anomala

Lindl. 1857, Journ. Linn. Soc. 1: 394. – H. grandiflora Ridl. 1923, Journ. Mal. Br.

Roy. As. Soc. 1, 1: 98. – H. rotundiloba J.J. Smith, 1926, Svensk Bot. Tidsskr. 20:

470. – H. hainanensis T. Tang et F.T. Wang, 1974, Act. Phytotax. Sin. 1, 1: 34. –

H. biloba (Ridl.) Seidenf. et Wood, 1992, Orch. Pen. Mal. Sing.: 95, fig. 38; Aver. et

Averyanova, 2003, l.c.: 40. – Zeuxine biloba Ridl. 1909, Journ. Fed. Mal. St. Mus. 4: 73.

Described from NE India (“Assam: Tangree”). Type (“Griffith s.n.”) – K (holotype).

Terrestrial or lithophytic herb up to 35 cm tall, with 3–5 narrowly-ovate leaves

5–8 cm long. Scape to 20 cm long, shortly hairy. Rachis white hairy, to 7 cm long, with

4–20 flowers. Bracts ovate, 5–7 mm long, shorter than ovary, acuminate. Sepals oblong,

5–6 mm long, olive-brown with white and red apex, lateral sepals spreading. Petals


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129

lanceolate, 5 mm long, white with reddish-brown base, connivent to median sepal. Lip

white with saccate greenish-pink base containing 2 short gland-like keels on each

side, mesochile short, channeled with inrolled edges, epichile with 2 recurved broadly

obovate lobes, each lobes 3–4.5 mm long, 2.5 mm wide. Column with short narrow

wings and prominent rostellum arms. Fig. 34, m, n; 37, d.

Ecology. Wet primary and secondary closed forests, often with bamboo, on

any soils. 800–1300 m. Fl. January – April, August. Not rare (VU).

Distribution: Vietnam (Dac Lac, Hoa Binh, Lao Cai, Quang Binh, So La),

NE India, Burma, Laos, S China, Malaya, Sumatra, Borneo, Sulawesi.

Studied specimens. Dac Lac, Chu Yang Sinh mt., 1996, Vu Ngoc Long s.n. (LE);

Hoa Binh, Da Bac, Phu Canh mt., HAL 396 (HN, LE); Lao Cai, Van Ban, HAL 2287, DKH 6987

(HN, LE); Quang Binh, Bo Trach, HAL 6060 (HN, LE); Quang Binh, Phong Nha – Ke Bang

national park, HAL 6241 (HN, LE); So La, Yen Chau, DKH 7225 (HN, LE, MO).

2. H. alta Ridl. 1896, Journ. Linn. Soc. Bot. 32:404.

Described from Malacca Peninsular (“Perak: Hermitage Hills, Kuala Kangsa

Valley”). Type (“Ridley s.n.”) – ?

Terrestrial herb up to 50 cm tall, at the base with 4–6 ovate to elliptic leaves

6–9 cm long distant on 2–3 cm; sheaths to 2 cm long. Scape to 20 cm long, shortly

hairy, with many hardly opening flowers. Floral bracts cuneate, hairy, 5–7 mm long,

shorter than ovary. Sepals yellow-pink, ovate, 5–6 mm long, hairy. Petals white, ovate

of the same length, connivent to median sepal. Lip 4–5 mm long, at the base, with 2

close irregularly lobed papillae on each side, epichile in form of small slightly reflexed

cordate tip less than 0.5 mm long. Column with tall spathulate wings equal or slightly

exceeding rostellum arms. Fig. 34, o-q; 37, e.

Distribution: Vietnam (Thua Thien – Hue). Peninsular Thailand, Kalimantan.

Ecology. Wet closed primary and secondary forests on silicate soils. 300–

400 m. Fl. March – April. Very rare (DD).

Studied specimens. Thua Thien – Hue Prov. Nam Dong HAL 7156 (HN, LE).

3. H. oblongifolia (Blume) Blume, 1858, Coll. Orch. Arch. Ind. Jap.: 102,

tab. 32; Seidenf. 1992, Orch. Indochina: 39; Aver. 1994, Identif. Guide Vietnam. Orch.:

32; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 40. – Etaeria

oblongifolia Blume, 1825, Bijdr. 6, fig. 14; id. 1825, ibid. 8: 410.

Described from Java (“Solassie Provinciae Tjanjor”). Type – L (BO, P)?

Terrestrial herb up to 40 cm tall, at the base with 4–7 shining narrowly-ovate

leaves 4–6 cm long. Scape shortly hairy, to 16 cm long with many hardly opening

flowers. Floral bracts narrowly-cuneate, 3–5 mm long, shorter than ovary. Sepals light

yellowish, ovate, 3–3.5 mm long, hairy. Petals white, oblique narrowly-obovate, 3–

3.5 mm long, connivent to median sepal. Lip 3–3.5 mm long, yellowish, with about 5–

6 slender papillae on each side, epichile brightly yellow, triangular, fat, finely papillose,

apiculate. Column at front with strongly sigmoid, triangular-acute wings, shorter than

rostellar arms. Fig. 35, a-c; 37, f.


130

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 34. Goodyera foliosa: a – flower, lip section and petals; G. schlechtendaliana: b – flower,

lip section and petals; G. arisanensis: c – flowering plant, d – flattened tepals and lip section,

side view; G. rhombodoides: e – flowering plant and leaf, f – flattened tepals, g – flattened lip

and lip section, side view, h – column, dorsal and frontal views; Herpysma longicaulis: i –

flowering plant, j – flattened sepals and flattened petals, k – lip, side view, l – column, frontal

and half-side views; Hetaeria anomala: m – flower, n – flattened tepals, column and lip;

H. alta: o – flattened tepals and column, p – lip, frontal view, q – column, side view.


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131

Distribution: Vietnam (Vungtau – Condao), Andaman islands, Thailand,

Malacca, Bonin, Ryukyu islands, Philippines, Indonesia, New Guinea, Australia, Pacific

islands.

Ecology. Wet primary and secondary closed forests on silicate soils. 200–

400 m. Fl. March – May. Locally common (VU).

Studied specimens. Vungtau – Condao, Con Dao, 1987, Averyanov 817, 840, 905

(HN, LE); 1989, Averyanov 502 (HN, LE).

4. H. affinis (Griff.) Seidenf. et Ormerod, 2001, Oasis, Suppl. 2: 9. – Goodyera

affinis Griff. 1851, Not. Pl. Asiat. 3: 391; id. 1851, Ic. Pl. Asiat. 3, tab. 351B, fig. 7. –

Cerochilus rubens Lindl. 1854, Gard. Chron. ser. 1: 87. – H. rubens (Lindl.) Benth.

ex Hook. f. 1890, Fl. Brit. Ind. 6: 115; Seidenf. 1992, Orch. Indochina: 39; Aver. 1994,

Identif. Guide Vietnam. Orch.: 33; Aver. et Averyanova, 2003, Updated Checklist

Orch. Viet.: 40.

Described from NE India (“Assam, n. Negrogam”). Type (“18-01-1836,

Griffith in Assam Herb. no. 315”) lost (Seidenf. & Ormerod, 2001, l.c.).

Terrestrial herb up to 45 cm tall, with 4–8 narrowly-ovate to ovate leaves 5–

10 cm long. Scape hairy, to 25 cm long, with many hardly opening flowers. Floral

bracts cuneate, sparsely hairy, 4–7 mm long, shorter than ovary. Pedicel and ovary

straight, densely adpressed to the rachis. Sepals greenish-pink, ovate, 4–5 mm long,

sparsely hairy. Petals white, oblique narrowly-obovate, hammer-like, with a semicircular

widening at upper edge faced to the dorsal sepal, 4–5 mm long, connivent to median

sepal. Lip 4–5 mm long, yellow, with 1–2 simple or irregular lobed fat papillae on each

side; epichile in form of very small slightly reflexed conduplicate, cordate white tip.

Column 1.5 mm tall, at front with small triangular inward turned wings near the apex,

shorter than small triangular acute rostellar arms. Fig. 35, d, e; 37, g.

Distribution: Vietnam (Dak Nong, Nghe An, Ninh Binh, Quang Tri, Thua

Thien – Hue, Tuyen Quang), Bhutan, NE India, Burma, China, Thailand, Vietnam.

Studied specimens. Dak Nong, Dak Glong, Ta Dung mt., HLF 5608 (HN, LE);

Nghe An, Con Cuong, Phu Pha May mt., HLF 3116 (HN, LE); Nghe An, Tuong Duong, HLF

6591 (HN, LE); Ninh Binh, Cuc Phuong national park, Henk van der Werff et al. 14209 (HN, LE,

MO); Quang Tri, Da Krong Nature Reserve, HLF 6256, HN, LE, d-EXSICCATES OF

VIETNAMESE FLORA 0064/HLF6256; Thua Thien – Hue, Nam Dong, HAL 10934 (HN, LE);

Tuyen Quang, Na Hang, HAL 220 (HN, LE).

5. H. nitida Ridl. 1896, Journ. Linn. Soc. Bot. 32: 404; Seidenf. Wood, 1992,

Orch. Pen. Mal. Sing.: 95, fig. 37 g-i, pl. 2, d.

Described from Malacca Peninsular (“Pulau Butong, Penang”). Type (“Curtis

s.n.”) – SING (holotype).

Terrestrial herb up to up to 45 cm tall, with 5–7 narrowly-ovate to ovate leaves

7–10 cm long. Scape hairy, to 25 cm long, with many hardly opening flowers. Floral

bracts cuneate, hairy, 5–8 mm long, shorter than ovary. Pedicel and ovary straight,

densely adpressed to the rachis. Sepals olive to pink, ovate, 5–6 mm long, hairy. Petals


132

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

white, oblique ovate, broadly-ovate to broadly-obovate, 5–6 mm long, connivent to

median sepal. Lip 5–5.5 mm long, yellow, with 2(3) irregular lobed fat papillae on each

side; epichile triangular, conduplicate, with fat edges and shortly apiculate tip, brightly

yellow. Column 1.5 mm tall, at front with small rectangular inward turned wings near

the apex, shorter than small triangular acute rostellar arms. Fig. 35, f, g; 37, e, i.

Distribution: Vietnam (Ha Giang, Quang Tri), Thailand, Malaya, Singapore.

Ecology. Wet closed primary forests on rocky limestone. 150–1200 m. Fl.

March – May. Rare (VU).

Studied specimens. Ha Giang, Quan Ba, Bat Dai Son mt., NTH 5420G (HN, LE);

Quang Tri, Da Krong Nature Reserve, HLF 6255, HN, LE, d-EXSICCATES OF VIETNAMESE

FLORA 0065/HLF6255.

Ludisia A. Rich. 1825, in Bory de St.-Vincent, Dict. Class. Hist. Nat. 7: 437.

Type: L. discolor (Ker-Gawl.) A. Rich. (Goodyera discolor Ker-Gawl.)

Lithophytic and occasionally epiphytic sympodial herb with fleshy succulent

creeping rhizome and ascending short leafy stem with 2–6 petiolate leaves and terminal

erect leafless inflorescence. Petiole and sheath 1–3 cm long; leaf blade ovate to broadelliptic,

4–10 cm long, 1.5–4 cm wide, dark velvety green to almost black, with 1–7

yellow or pink longitudinal veins, occasionally reticulated with veins network. Scape

10–18 cm tall, with 2–3 pale pink, hyaline bracts. Rachis 2–14 cm long, hairy, with 1–

12 laxy flowers. Floral bracts cuneate, 1–1.5 cm long, a little shorter than ovary.

Flowers resupinate, widely open, 1.2–1.7 cm across, white, with yellow anther. Sepals

ovate, 10 mm long. Petals narrowly-ovate of same length, connivent to dorsal sepal

forming hood. Lip twisted, with short 2-lobed spur; spur inside with 2-lobed sessile

gland on each side; lip blade 7–9 mm long, clawed, claw channeled, epichile 2-lobed,

with spreading oblong lobes. Column twisted, stigma single on front of column. Fruit

ellipsoid capsule, 1.5 cm long, 4–5 mm wide. Pollinia 2, sectile, with caudicle and

viscidium. Fig. 35, h-k; 43, a-c.

Monotypic genus.

1. L. discolor (Ker-Gawl.) A. Rich. 1825, in Bory de St.-Vincent, Dict. Class.

Hist. Nat. 7: 437; Seidenf. 1992, Orch. Indochina: 29; Aver. 1994, Identif. Guide Vietnam.

Orch.: 41; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 43. – Goodyera

discolor Ker-Gаwl. 1818, Bot. Reg. 4, tab. 271. – Anoectochilus dawsonianus

H. Low ex Reichenb. f. 1868, Gard. Chron. (1868): 1038. – Haemaria otletae Rolfe,

1891, Ill. Hort. 38: 31, tab. 524. – H. pauciflora Gagnep. 1933, Bull. Soc. Bot. France, 80:

349. – L. dawsoniana (H. Low ex Reichenb. f.) Aver. 1988, Bot. Journ. (Leningrad), 73,

3: 432. – L. otletae (Rolfe) Aver. 1988, Bot. Journ. (Leningrad), 73, 3: 432.

Described from S China (“from the Brasils” – sphalm., sec. Lindl., 1840, Gen.

Sp. Orch. Pl.: 490). Lectotype: Icon “Goodyera discolor” (Bot. Reg. 4, t. 271 –

Ormerod, 2002, Lindleyana 17, 4: 211).

Ecology. Broad-leaved forests and shrubs on any kind of rocks, usually along

rocky river valleys. 50–1250 m. Fl. February – April. Not rare (LR).


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133

Distribution. Vietnam (probably all provinces), Myanmar, S China, Thailand,

Laos, Cambodia, Malacca Peninsular, Sumatra.

Studied specimens. Dac Lac, 24 Dec. 1979, Bien 1167 (HN, LE); Dak Lac, Chu

Yang Sinh mt., VH 5935 (HN), VH 6014 (HN); Ha Giang, Quan Ba, Bat Dai Son mt., NTH 5420D

(HN, LE); Ha Tinh, Huong Son, HAL 1340 (HN), VA 12 (HN, MO), VA 429 (HN, MO), VA 858

(HN, NO); Kontum, Sa Thay, Chu Mom Ray, P-7906b (HN); Nghe An, Tuong Duong, HLF

6570 (HN, LE), HLF 6720 (HN); Ninh Binh, Cuc Phuong national park, 1971, Bao 471, CPNP

5148 (HN), NMC 706 (HN), NTH 4104 (HN), MVX 36 (HN), DDS 11761 (HN), HAL 1605 (HN),

HAL 1655 (HN), sine date, Bao 471 (HN), sine date, MVX 172 (HN). Ninh Thuan, Ninh Hai,

HLF 4015 (HN); Ninh Thuan, Ninh Hai, Nui Chua mt., HLF 4281 (HN, LE); Phu Khanh, Nha

Trang, 11 June 1989, Averyanov, Kudryavtzeva 201 (HN, LE); Quang Binh, Bo Trach, HAL

6078 (HN, LE), HAL 6279 (HN), HAL 6350 (HN, LE), HAL 6351 (HN), d-EXSICCATES OF

VIETNAMESE FLORA 0009/HAL6351; Quang Binh, Minh Hoa, HAL 5865 (HN, LE), HAL

5926 (HN), HAL 5983 (HN); Quang Binh, Minh Hoa, Ke Bang, VH 4771 (HN, LE); Quang Binh,

Quang Ninh, HAL 6310 (HN); Quang Tri, Da Krong, HLF 6217 (HN); Quang Tri, Huong Hoa,

HLF 5933 (HN, LE); Quangnam – Danang, Cu Lao Cham, 23 March 1987, Averyanov,

Kudryavtzeva 541 (HN, LE); Thanh Hoa, Ba Thuoc, HAL 1076 (HN); Thanh Hoa, Thach

Thanh, HAL 2891 (HN); Thua Thien – Hue, A Luoi, HAL 7994 (HN); Thua Thien – Hue, Bach

Ma national park, HLF 1669 (HN, LE), HLF 1818 (HN, LE), HLF 2695 (HN), HLF 2696 (HN);

Thua Thien – Hue, Huong Thuy, HAL 8111 (HN); Vinh Phu, Tam Dao, 1988, Averyanov et al.

LX-VN s.n. (LE), LX-VN 2383 (HN, LE); Vungtau – Condao, Con Dao, 23 March 1989, Averyanov,

Kudryavtzeva 307 (HN, LE).

Note. Very variable species, particularly on coloration of leaves. Widespread

common orchid in lowland areas of Vietnam. It is widely collected for traditional

oriental medicine and as ornamental plant for cultivation known under popular name

“jewel orchids”.

Macodes Lindl. 1840, Gen. Sp. Orch. Pl.: 496.

Type: M. petola (Blume) Lindl. (Neottia petola Blume)

Terrestrial sympodial herbs with creeping succulent rhizome rooting at nodes,

ascending erect leafy floriferous stem and terminal few- to many-flowered raceme.

Leaves few, spirally arranged at the base of stem, shortly petiolate, sheathing at the

base, glabrous, convolute, asymmetric, acute, bright green with fine network of yellowgolden

nerves. Scape leafless, with 2–4 sterile bracts. Flowers small, not resupinate.

Sepals free; petals connivent to median sepal forming hood. Lip spurless, twisted, 3lobed.

Hypochile deeply saccate with 2 fat glands on lateral walls. Sidelobes triangular,

acute. Epichile spathulate, with narrow stalk-like claw widening into more or less

reflexed circular apex. Column short, twisted, on front with 2 thin close parallel rostellar

arms descending downwards into saccate hypochile and 2 hemicircular wings. Anther

acute. Stigma large, not divided.

10(1) species. Ryukyu Islands, Peninsular Malacca, Indonesia to Philippines,

New Guinea and Vanuatu.

1. M. cupida Ormerod, 2002, Lindleyana, 17, 4: 212.


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Described from S Vietnam (“Dalat, cult. Hort. Bot. Parisiensis, 1959”). Type

(“Tixier M-918”) – P.

Rachis pubescent, laxy 4–5 flowered. Floral bracts ovate-elliptic, acute,

pubescent, 9.5 –9 mm long, 4 mm wide. Pedicel and ovary pubescent, to 12 mm long.

Flowers glabrous, not resupinate. Sepals narrowly-ovate, acute, 8 mm long, 4 mm

wide; lateral sepals recurved, oblique. Petals oblique narrowly-ovate, 8 mm long, 3

mm wide. Lip 3-lobed, 8.5 long, 5 mm wide. Hypochile subquadrate, saccate, sidelobes

broadly rounded, with the apical quarter membranaceous; inside along median vein

hypochile with a low fleshy keel terminating apically in an arrow shape about 4 mm

long, 5 mm wide; hypochile glands terete with an enlarged apex. Claw linear, 4 mm

long. Epichile cordate, acute, 1.5 mm long, 2 mm wide. Column erect, 6.5 mm tall.

Anther basally ovoid, long rostrate, 5 mm long. Rostellar arms linear, 4 mm long.

Stigma ventral, at the base of rostellum. Column wings subquadrate, ventral, 1 mm

across. Fig. 35, l-n.

Distribution. Vietnam (Lam Dong?).

Note. Insufficiently known species described on the base of fragment of

inflorescence collected from alone cultivated specimen, origin of which needs

verification. It may be related to widely cultivated M. petola, from which differs in

broader narrowly-ovate petals and twice longer anther and rostellum.

Myrmechis Blume, 1859, Coll. Orchid.: 76

Type: M. gracilis (Blume) Blume (Anoectochilus gracilis Blume)

Very small terrestrial, lithophytic or occasionally epiphytic sympodial herbs

with creeping slender rhizome, ascending erect leafy floriferous stem and 1–2(3)

terminal flowers. Leaves few, spirally arranged, petiolate, sheathing at the base,

glabrous, convolute, uniformly green, distant or crowded near apex. Scape short, with

1–3 sterile bracts. Flowers small, sessile, resupinate, hardly opening. Sepals free,

subsimilar, lateral concave, oblique. Petals connivent to median sepal forming hood.

Lip connate to the base of the column, long-clawed. Hypochile concave, with short,

thick, paired warts on lateral sides. Mesochile prominent, narrow, with incurved margin.

Epichile recurved, broad, 2-lobed. Column short, erect, with short erect rostellum and

2 small conical wings at front. Anther broadly-ovate. Pollinia 2, sectile, with caudicle

and viscidium. Stigmas 2 on short lateral projections.

15(1) species. Tropical and subtropical Asia.

1. M. pumila (Hook. f.) T. Tang et F.T. Wang, 1951, Acta Phytotax. Sin. 1:

69. – Odontochilus pumilus Hook. f. 1890, Fl. Brit. India, 6: 99. – Zeuxine

langbianensis N.T. Tich, 2001, Orch. Vietnam, 1: 214.

Described from NE India (“India, Sikkim: alt. 8.000–10.000 ft”). Type (“J.D.Hooker

325A”) – K.

Terrestrial herb 6–10 cm tall. Leaves shortly petiolate, broadly-ovate to almost

circular, to 1 cm long and wide, uniform green. Scape 2.5–3 cm long, with 1–2 flowers.

Floral bracts ovate to broadly-ovate, 4.5 mm long, 3.8 mm wide, ciliate along margin.

Sepals sub-similar, ovate 6 mm long, obtuse. Petals oblique narrowly-ovate, suddenly


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Fig. 35. Hetaeria oblongifolia: a – flattened tepals and lip, b – column and lip, side view, c –

column, frontal view; H. affinis: d – flattened tepals and lip, e – column, frontal view; H. nitida:

f – flattened tepals and lip, g – column, frontal view and section of spur base. Ludisia discolor:

h – flowering plant, i – flattened tepals and lip, j – flower, k – column, side view and lip;

Macodes cupida: l – flattened sepals and petal, m – flattened lip, n – column, side view;

Myrmechis pumila: o – flowering plant, p – flattened sepals, petal and lip, q – column and lip,

side view, r – floral bract.


136

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

narrowed near the apex, pyriform, 5.8 mm long, 2.2 mm wide. Lip 5.6 mm long.

Hypochile saccate, with 1 prominent midvein and 1 bilobed gland on each lateral vein.

Epichile 2-lobed, with spreading oblique obovate lobes. Column short, with small hornlike

frontal wings. Fig. 35, o-r.

Ecology. Wet broad-leaved closed mossy forests on silicate soils. 2100–2150m.

Fl. July. Very rare (EN).

Distribution. Vietnam (Kon Tum, Lam Dong), Bhutan, Nepal, NE India,

Myanmar, S China, Thailand.

Studied specimens. Kon Tum, Ngoc Linh mt., VH 627a (specimen lost); Lam

Dong, Lac Duong, Langbian mt., T.20.07.94b (icon “Zeuxine langbianensis” N.T. Tich, 2001,

l.c.: 215).

Odontochilus Blume, 1859, Coll. Orchid.: 79. – Evrardià Gàgnep. 1932, Bull.

Mus. Hist. Nat. (Paris) 2 ser. 4, 5: 596. – Pristiglottis Cretz. et J.J. Smith, 1934, Acta

Fauna Fl. Universali, ser. 2, Bot. 1, 14: 4. – Evrardianthe Ràuschert, 1983, Feddes

Repert. 94: 433. – Evrardianà Aver. 1988, Bot. Journ. (Leningràd) 73, 3: 432.

Lectotype: O. flavescens (Blume) Blume (Anoectochilus flavescens Blume)

Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome

rooting at nodes, ascending erect leafy floriferous stem and terminal lax, few- to many

flowered spike. Leaves few, spirally arranged, petiolate, sheathing at the base, glabrous,

convolute, asymmetric, uniformly green. Scape leafless, with 2–4 sterile bracts.

Flowers, sessile, resupinate or not. Sepals free. Petals very oblique, connivent to median

sepal forming hood, lateral sepals spreading, concave, enclosing saccate base of lip.

Lip adnate to column base. Hypochile concave to saccate to globose, with 2 fleshy

branched papillae on lateral sides, rare without glands. Mesochile channeled, long to

shortly fimbriate-toothed. Epichile with 2 divergent flabellate spreading lobes. Column

short, fleshy, winged or keeled, with 2 rostellar arms. Anther triangular-ovate, acute.

Pollinia 2, sectile, with stipe and viscidium. Stigma ventral, 1 or 2.

40(9) species. Tropical mainland Asia, Indonesia, New Guinea, Pacific Islands

to Polynesia and adjacent islands.

Key to species

1. Plants achlorophyllous, yellowish to pink-brown, all leaves reduced into small

yellowish bracts .................................................................................................................. 2

– Plants with normally developed green or purple-brown leaves ..................................... 3

2. Flowers not resupinate, yellow to red-orange; lip 12–15 mm long; epichile lobes lacinate

................................................................................................................................ 1. O. poilanei

– Flowers resupinate, with brown sepals and white lip; lip 9–10 mm long; epichile lobes

with entire margin ...................................................................................... 2. O. saprophyticus

3. Leaves purple-brown to black-violet; scape with 1–2(3) not resupinate flowers;

mesochile with purple-brown flanges ................................................................. 3. O. elwesii

– Leaves green; scape normally more than with 3 resupinate flowers; lip pure white or

yellow .................................................................................................................................... 4


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Fig. 36. Cheirostylis latipetala: a (HAL 8526 and DKH 7659c, type); C. spathulata: b (Thailand,

s. coll.); Erythrodes hirsuta: c (HAL 2266 and HLF 5755); E. blumei: d (Vietnam, s. loc.);

Goodyera procera: e (HAL 6924 and HLF 5756); G. fumata: f (LX-VN 1811); G. hispida: g (HAL

9385); G. biflora: h (HAL 11135); G. viridiflora: i (DKH 7400 and HAL 6057).


138

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 37. Goodyera foliosa: a (HAL 3967 and HAL 10027); G. schlechtendaliana: b (HLF 5257),

c (HAL 8404); Hetaeria anomala: d (HAL 6241); H. alta: e (HAL 7156); Hetaeria oblongifolia:

f (Averyanov 817, 1987); H. affinis: g (HLF 6591); H. nitida: h (HLF 6255 and NTH 5420G), i

(HLF 6255).


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4. Lip at the base straight, not concave or hardly concave, without any warts or glands

inside .............................................................................................................. 4. O. acalcaratus

– Lip at the base distinctly concave to saccate, with distinct lateral warts or glands

inside ..................................................................................................................................... 5

5. Ovary glabrous; column with 2 stigmas ........................................................................... 6

– Ovary tomentose or hairy; column with 1 stigma ............................................................ 7

6. Lip white, with convergent semi-circular to broadly-obovate lobes, slightly erose

along margin, 8–9 mm long; spike few-flowered, 2–4 cm long; hypochile glands more

or less straight, not uncinnate ........................................................................ 5. O. brevistylus

– Lip yellow, with rather triangular-flabellate lobes, divided by broad sinus, entire along

margin, 9–12 mm long; spike many-flowered, 4–12 cm long; hypochile glands uncinnate

.......................................................................................................................... 6. O. lanceolatus

7. Ovary and sepals densely hairy; lip 10–12 mm long; flanges of mesochile entire or

with few low insignificant teeth; lateral sepals not spreading, about 8 mm long; epichile

lobes 6 mm long .................................................................................................. 7. O. umbrosus

– Ovary and sepals sparsely short hairy; lip 16–20 mm long; flanges of mesochile with

3–5 long fringes; lateral sepals, 8–10 mm long; epichile lobes 8–12 mm long ............. 8

8. Sepals and petals uniform greenish, with 1–3 longitudinal veins, without distinct

pinnate venation from midvein ............................................................................. 8. O. tortus

– Sepals and petals brownish to brownish-pink, with unclear whitish pinnate tessellation

and distinct pinnate venation from mid-vein ..................................................... 9. O. inabai

1. O. poilanei (Gagnep.) Ormerod, 2002, Lindleyana, 17, 4: 225. – Evrardia

poilanei Gagnep. 1932, Bull. Mus. Nat. Hist. (Paris) 2, ser. 4, 5: 596. – Hetaeria

poilanei (Gagnep.) T. Tang et F.T. Wang, 1951, Acta Phytotax. Sin. 1: 71. – Evrardianthe

poilanei (Gagnep.) Rаuschert, 1983, Feddes Repert. 94: 433; Seidenf. 1992,

Opera Bot. 114: 34; Aver. 1994, Identif. Guide Vietnam. Orch.: 46. – Evrardianа

poilanei (Gagnep.) Aver. 1988, Bot. Journ. (Leningrаd) 73, 3: 432. – Chamaegastrodia

poilanei (Gagnep.) Seidenf. et A.N. Rao, 1994, Nord. Journ. Bot. 14: 297; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 20.

Described from southern Vietnam (“Annam, Dаlаt”). Type (“Evrаrd, 1807”)– P.

Terrestrial achlorophyllous leafless herb up to 20 cm tall. Stem yellowish, glabrous,

with many yellowish, broad, obtuse bracts and terminal lax spike with 2–10

flowers. Rachis shortly pubescent. Floral bracts ovate, acute, 8–10 mm long, glabrous,

ciliate along margin. Ovary shortly hairy, 8–10 mm long. Flowers not resupinate, orange.

Sepals shortly hairy at the base; median sepal ovate 4–6 mm long; lateral sepals

oblique-oblong, 8–10 mm long. Petals, lanceolate, oblique, obtuse, as long as median

sepal. Lip 12–18 mm long. Hypochile saccate, with 2 small simple warts. Mesochile

narrow, with pectinate irregular toothed, finely papillose flanges. Epichile lobes

spreading, triangular-flabellate, 6–9 mm long, 5–7 mm wide, irregular toothed or laciniate,

with prominently elongated 2 adaxial lobules. Column stalked, 2–3 mm tall, with 1 U–

shaped stigma, 2 long rostellar arms and 2 large parallel wings at front below stigma.

Anther ovate, acute, 1.6–2 mm long. Fig. 38, a-c.

Ecology. Primary and secondary humid broad-leaved forests on silicate soils.

1500–1600 m. Fl. October – November. Very rare (EN).


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Distribution. Vietnam (Kon Tum, Lam Dong), Myanmar, Thailand, Japan,

SChina.

Studied specimens. Kontum, mt. Mamray, Poilane 18204 (P); Lam Dong, Lac

Duong, WP 1212B (HN, LE).

2. O. saprophyticus (Aver.) Ormerod, 2003, Taiwania, 48, 3: 141. –

Pristiglottis saprophytica Aver. 2003, in Aver. et Averyanova, Updated Checklist

Orch. Viet.: 54, 90, fig. 9.

Described from southern Vietnam (“Lam Dong Prov., Lac Duong Distr., ...

Nui Gia Rich ridge ...”). Type (“VH 3977a”) – HN (holotype), LE (isotype).

Terrestrial achlorophyllous leafless herb up to 22 cm tall. Rhizome rootless,

densely covered with root hairs. Stem pink-brown, glabrous, with many hyaline

acuminate bracts and terminal lax spike with 3–15 flowers. Rachis pubescent to 10

cm long. Floral bracts elliptic, acuminate hairy, 6–10 mm long. Ovary pubescent, 7–9

mm long. Sepals sub-similar, narrowly-ovate, sparsely hairy, pink-brown to olive-brown,

5–6 mm long; lateral sepals slightly oblique, spreading. Petals white, lanceolate, obtuse,

as long as sepals. Lip white, 9–10 mm long. Hypochile saccate, with 2 large simple

glands. Mesochile narrow, 4–5 mm long, with numerous warts and papillae on flanges.

Epichile lobes spreading, sub-quadrate to hemi-circular, 4–4.5 mm long, 3.5–4 mm

wide. Column narrowing to the base, 3.5–4 mm tall, with 1 large circular stigma, 2

long rostellar arms and 2 parallel hemicircular wings at front. Anther narrowly-ovate,

acute, 2–2.5 mm long. Fig. 38, d-h; 43, d.

Ecology. Primary humid broad-leaved forests on silicate soils, commonly on

rich soils along shady stream slopes. 450–1500 m. Fl. April – May. Very rare (EN).

Distribution. Vietnam (Lam Dong, Thua Thien - Hue).

Studied specimens. Thua Thien – Hue, Huong Thuy, HAL 8194 (HN, LE); d-

EXSICCATES OF VIETNAMESE FLORA 0018/HAL8194.

3. O. elwesii C.B. Clarke ex Hook. f. 1890, Fl. Brit. Ind. 6: 100. – Anoectochilus

elwesii (C.B. Clarke ex Hook. f.) King et Pantl. 1898, Ann. Roy. Bot. Gard.

(Calcutta), 8: 297; Seidenf. 1992, Opera Bot. 114: 31; Aver. 1994, Identif. Guide Vietnam.

Orch.: 44; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 10. – A. daoensis

Gagnep. 1951, Bull. Mus. Natl. Hist. Nat. 2 ser. 22, 5: 632; Seidenf. 1992, l.c.: 30;

Aver. 1994, Identif. Guide Vietnam. Orch.: 44.

Described from NE India (“Darjeeling, Rishap; Khasia Hills, Shillong; Munipore,

Kohima, Naga Hills”). Syntypes (“Clarke 12251; Clarke 44609; Prain 17”) – K.

Terrestrial and lithophytic herb to 20 cm tall with 2–5 petiolate leaves at the

base. Leaves ovate, acute, purple-brown to black-violet, 1.5–4.5 cm long. Scape to 15

cm long, hairy, with 1–3 acute, hyaline bracts and 1–2(3) lax not resupinate flowers.

Floral bracts lanceolate, acuminate, hairy, to 12 mm long. Ovary 10–15 mm long,

densely pubescent. Sepals olive to brown-green, long hairy, with acute, strongly recurved

apex; median sepal ovate, 9–11 mm long, with light pinkish apex; lateral sepals obliqueovate,

10–12 mm long, with, concave, hemi-spherical base, embracing lip base. Petals


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141

white, hemi-circular, strongly oblique, broad at the middle, attenuate toward the apex,

as long as median sepal. Lip white, 1.6–2.5 cm long, basally globular, bifid, with 2

prominent conical glands. Mesochile pectinate with 4–6 purple-brown fringes. Epichile

lobes triangular-flabellate, irregular-undulate along margin, 1–1.5 cm long, 6–8 mm

wide, divided by a broad sinus. Column twisted, 2–3 mm tall, with 2 stigmas and 2

appendages at the base. Anther beaked. Fig. 38, i-o; 43, e.

Ecology. Primary and secondary wet broad-leaved forests, often with bamboo

on any kind of soils. 300–1200 m. Fl. May – July. Not common (VU).

Distribution. Vietnam (Bac Kan, Lai Chau, Lao Cai, Quang Binh, Thanh

Hoa, Tuyen Quang, Vinh Phuc), Bhutan, Nepal, NE India, Myanmar, Thailand, S China,

Taiwan.

Studied specimens. Bac Kan, Ba Be, HLF 612 (HN, LE); Bac Kan, Cho Don, HAL

4791 (HN, LE); Lai Chau, Sin Ho, HAL 8835 (HN, LE); Lao Cai, Van Ban, HAL 2003 (HN, LE);

Quang Binh, Minh Hoa, HAL 5957 (HN, LE); Thanh Hoa, Ba Thuoc, HAL 3372 (HN, LE);

Tuyen Quang, Na Hang, HAL 1720 (HN, LE); Vinh Phu, Tam Dao, 23 Jan. 1988, Averyanov et

al. LX-VN s.n. (LE), P-7452 (HN, LE).

4. O. acalcaratus (Aver.) Ormerod, 2002, Lindleyana, 17, 4: 225. – Anoectochilus

acalcaratus Aver. 1996, Bot. Journ. (St. Petersburg), 81, 10: 73, fig. 1; Aver.

et Averyanova, 2003, Updated Checklist Orch. Viet.: 10.

Described from southern Vietnam (“Prov. Kontum ... N slope of Ngoc Linh

mountain ...”). Type (“VH 627”) – LE (holotype), AAU, HN, MO, P (isotypes).

Terrestrial herb up to 20 cm tall with 3–5 dark velvety green leaves at the

base. Leaves shortly petiolate, narrowly-ovate to ovate, 3.5–7 cm long, usually finely

undulate along margin. Scape 3–6 cm tall with 2–3 small sessile leaves. Rachis hairy

2–5 cm long with 4–8 lax flowers. Floral bracts hairy, 8–10 mm long, longer than

ovary. Sepals sub-similar, light green, elliptic, sparsely hairy, 8–10 mm long 3–4 mm

wide. Petals white, lanceolate, 6 mm long. Lip white, little longer than sepals, at the

base straight, not concave, without glands, with longitudinal low keel. Mesochile narrow,

straight, 5 mm long, with 3–5 small teeth. Lobes of epichile spreading, hemicircular, 4–

4.5 mm long, 2.5–3 mm wide. Column 3 mm tall, with 2 stigmas. Fig. 38, p-t.

Ecology. Primary evergreen mossy forests on silicate soils, commonly along

shady wet streams canyons. 2100–2450 m. Fl. June – July. Locally common (VU).

Distribution. Vietnam (Kon Tum).

Studied specimens. Kontum, Ngoc Linh mt., VH 539 (HN, LE).

5. O. brevistylus Hook. f. 1890, Fl. Brit. Ind. 6: 100. – Anoectochilus brevistylus

(Hook. f.) Ridl. 1907, Mat. Fl. Malay. Penins. 1: 214; Seidenf. 1992, Opera

Bot. 114: 31; Aver. 1994, Identif. Guide Vietnam. Orch.: 43; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 10. – Anoectochilus tonkinensis Gagnep. 1931,

Bull. Mus. Paris 2, ser. 3, 7: 679.

Described from Malacca Peninsula (“Perak, on Gunong Batu Pateh, alt. 3400

ft.”). Type (“Wray s.n.”) – K?


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 38. Odontochilus poilanei: a – flowering plant, b – flattened tepals and lip, c – column,

side view; O. saprophyticus: d – flowering plant, e – flattened tepals and lip, f – fragment of

mesochile, g – column, side view, h – pollinarium; O. elwesii: i – inflorescence, j – flower and

flower without lateral sepals and epichile, k – section of spur and hypochile, l – flattened lip,

m– section of spur base, n – column, frontal view, o – pollinarium; O. acalcaratus: p – flowering

plant, q – flower and fragment of inflorescence, r – flower without lateral sepal and column with

lip, side view, s – flattened tepals and lip, t – column, ventral and dorsal views.


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Herb to 30 cm tall with 5–6 distant pure green petiolate leaves. Leaves narrowly-ovate

to ovate, acute, 4.5–7 cm long. Scape short, 2–4 cm long, with lax spike

2–4 cm long of 6–10 flowers. Floral bracts narrowly-ovate, acuminate, to 10 mm long,

a little shorter than ovary. Ovary glabrous. Sepals sub-similar, ovate, 7–8 mm long,

green, sparsely hairy; lateral sepals swollen at the base, enclosing base of lip. Petals,

strongly oblique, falcate with attenuate apex, as long as median sepal. Lip white, 10–

14 mm long, basally shortly globular, bifid, with prominent longitudinal ridge and flat

quadrangular gland on each side. Mesochile 7–8 mm long, laterally with 3–4 long

fringes. Epichile lobes 8–9 mm long, 5–6 mm wide, semi-circular to broadly-obovate,

slightly irregularly toothed, divided by broad sinus. Column short, 2.5–3 mm tall, with 2

stigmas, short rostellar arms and 2 small insignificant wings at the base. Fig. 39, a, b.

Ecology. Humid broad-leaved forests. 1400–1600 m. Fl.? Very rare (DD).

Distribution. Vietnam (Lao Cai), Thailand, S China, Taiwan, Malacca Peninsula.

Studied specimens. Chapa, Tonkin, Petelot s.n. (P).

6. O. lanceolatus (Lindl.) Blume, 1859, Coll. Orchid.: 80. – Anoectochilus

lanceolatus Lindl. 1840, Gen. Sp. Orch. Pl.: 499; Seidenf. 1992, Opera Bot. 114: 30;

Aver. 1994, Identif. Guide Vietnam. Orch.: 43; Aver. et Averyanova, 2003, Updated

Checklist Orch. Viet.: 11.

Described from NE India (“Assam; Khasia, Mishmee Hills”). Syntypes (Mack

s.n.; Griffith s.n.) – K.

Terrestrial and lithophytic herb to 35 cm tall with 5–7 distant petiolate leaves.

Leaves ovate, 4–10 cm long. Scape short, 5–6 cm long, pubescent with lax many

flowered spike 4–10 cm long. Floral bracts narrowly-ovate acuminate, 8–15 mm long.

Ovary 8–10 mm long, glabrous. Sepals and petals pale green; median sepal ovate,

acute, 5–7 mm long; lateral sepals ovate-oblong, oblique, spreading, 7–8 mm long.

Petals strongly oblique, hemi-circular with attenuate apex, 5–7 mm long, 2–2.5 mm

wide. Lip yellow, 16–18 mm long, basally shortly globular, bifid, with 2 slender uncinnate

glands. Mesochile 6–8 mm long, laterally with 5–7 rair fringes. Epichile lobes spreading,

triangular-flabellate, 8–10 mm long, 4–6 mm wide. Column stout, 2–3 mm tall, with 2

stigmas, large elongate, bifid rostellum and 2 prominent vertical parallel wings at the

base. Anther narrowly-ovate, beaked, 2.5–3 mm long. Fruit erect fusiform capsule 1–

1.2 cm long. Fig. 39, c–e.

Ecology. Primary humid broad-leaved forests, usually along shady stream

canyons on any kind of soils. 1200–1300 m. Fl. August – September. Rare (EN).

Distribution. Vietnam (Ha Giang, Ha Noi, Lao Cai), Bhutan, Nepal, NE

India, S China, Thailand.

Studied specimens. Bavi mt., Balansa 2005 (P); Chapa, Petelot 5194 (P); Ha

Giang, Vi Xuyen, DKH 5316 (HN, LE, MO).

7. O. umbrosus (Aver.) Ormerod, 2002, Lindleyana, 17, 4: 225. – Pristiglottis

umbrosa Aver. 1996, Bot. Journ. (St. Petersburg) 81, 10: 78, fig. 3; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 54.


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Described from southern Vietnam (“Kontum Prov., Dak Gley Distr., ...”).

Type (VH 1932) – LE (holotype), HN, MO (isotypes).

Terrestrial and lithophytic herb to 25 cm tall with 5–7 more or less distant

petiolate leaves. Leaves ovate, dark green, sometimes finely undulate along margin, to

8 cm long, 3.5 cm wide. Scape 5–7 cm long, usually with 1 rudimentary leaf. Rachis

pubescent, 3–8 cm long, with 8–14 lax flowers. Floral bracts pubescent, 8–10 mm

long, not longer than ovary. Ovary 10–12 mm long, densely pubescent. Sepals brown

to brown-green, sparsely pubescent; median sepal ovate, acute, 6–7 mm long; lateral

sepals ovate, oblique, 8 mm long. Petals white, strongly oblique, with broad base and

attenuate apex, 6 mm long, 3 mm wide at the base. Lip white, 10–12 mm long, basally

shortly globular, bilobulate, with 2–3 close glands on each side. Mesochile 6 mm long,

laterally with 4–5 short fringes. Epichile lobes spreading, ovate, 6 mm long, 4 mm

wide. Column twisted, about 1 mm long, with 1 stigma and 2 small vertical wings at

the base. Anther narrowly-ovate, brightly red, 4 mm long. Fig. 39, f-k; 43, f.

Ecology. Primary humid broad-leaved forests, usually along shady stream

canyons on silicate soils. 1000–1500 m. Fl. October – January. Rare (EN).

Distribution. Vietnam (Dak Nong, Kon Tum).

Studied specimens. Dak Nong, Dak Glong, HLF 5611a (HN, LE).

8. O. tortus King et Pantl., 1896, Journ. As. Soc. Bengal, 2, Nat. Hist. 65:

125. – Anoectochilus tortus (King et Pantl.) King et Pantl. 1898, Ann. Roy. Bot.

Gard. (Calcutta) 8: 298, tab. 396. – Pristiglottis torta (King et Pantl.) Aver. 1996,

Bot. Journ. (St. Petersburg) 81, 10: 78. – A. repens auct. non (Downie) Seidenf. et

Smitinand: Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 11.

Described from northern India (“Darjeeling (Duars) Kumai”). Type (Pantling

354) – CAL (holotype), BM, K (isotypes).

Terrestrial and lithophytic herb to 30 cm tall with 4–7 distant petiolate leaves.

Leaves ovate, uniform green, 4–7 cm long. Scape short, 4–5 cm long, pubescent with

lax many flowered spike 4–10 cm long. Floral bracts narrowly-ovate, acuminate, finely

pubescent, 8–12 mm long. Ovary hairy, 8–12 mm long. Sepals and petals greenish.

Sepals sparsely long hairy; median sepal ovate, acute, 5–8 mm long; lateral sepals

ovate-oblong, oblique, spreading, 7–10 mm long. Petals strongly oblique, falcate with

attenuate apex, as long as median sepal. Lip pure white, 12–16 mm long, basally

shortly globular, with 2 thick close massive straight conical glands on each side.

Mesochile 5–7 mm long, laterally with 5–7 pair long fringes. Epichile lobes spreading,

hemi-circular to broadly-obovate, 4–7 mm long and wide. Column stout, 2.5–3.5 mm

tall, with 1 semi-circular stigma, conical rostellar arms and 2 small vertical wings at

the base. Anther narrowly-ovate, beaked, 2.5–3 mm long. Fig. 39, l-n.

Ecology. Closed humid broad-leaved forests on rocky limestone in shady, wet

places. 400–950 m. Fl. June – August. Very rare (EN).

Distribution. Vietnam (Ninh Binh). Bhutan, N India, S China, Taiwan.

Studied specimens. Ninh Binh, Cuc Phuong national park, HAL 1670 (HN, LE).


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145

Fig. 39. Odontochilus brevistylus: a – flower, b – section of spur base; O. lanceolatus: c –

flower, d – section of spur base, e – column, side view; O. umbrosus: f – flowering plant, g –

flower, h – flattened tepals and lip, i – anther and base of spur, view from above, j – base of lip,

view from below, k – column, half-side and frontal views; O. tortus: l – flattened tepals, m –

flattened lip and column, side view, n – section ob spur base; O. inabai: o – flower.


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

9. O. inabai (Hayata) T.P. Lin, 1975, Nat. Orch. Taiwan: 216. – Anoectochilus

inabai Hayata, 1914, Icon. Pl. Formos. 4: 102, pl. 14.

Described from Taiwan (“Rinkiho, ad 5000 ped. alt.”). Type (H. Inaba, Aug.

1911) – TI?

On its vegetative features this species resembles O. tortus, but differs in

slightly larger flowers, as well as brownish to brownish-pink sepals and petals, with

irregular pinnate whitish tessellation and pinnate venation. Fig, 39, o; 43, g.

Ecology. Closed humid broad-leaved forests on rocky limestone. 900–1000 m.

Fl. June – August. Very rare (EN).

Distribution. Vietnam (Hoa Binh), Taiwan.

Studied specimens. Hoa Binh, Mai Chau, HAL 9002 (HN).

Orchipedum Breda, Kuhl et Hasselt, 1827, Gen. Sp. Orch. Asclep.: tab. 10.

Type: O. plantaginifolium Breda

Terrestrial sympodial herbs with creeping succulent rhizome rooting at nodes,

ascending erect leafy floriferous stem and terminal many flowered spike. Leaves

few, spirally arranged, petiolate, sheathing at the base, glabrous, convolute, asymmetric,

uniformly green. Scape leafless, with 2–4 large sterile bracts. Flowers, sessile,

resupinate. Sepals free. Petals oblique-obovate, joined in apical part and connivent to

median sepal forming hood; lateral sepals spreading, with concave base, enclosing

saccate base of lip. Lip adnate to column base, spurred, 3-lobed. Spur globose, slightly

bifid, curved forwards under the blade, containing dense group of fleshy hairs on each

side. Base of lip with erect rounded side lobes embracing column base, inside with 2

erect semi-circular parallel keels. Median lobe with narrow clawed base suddenly

broadening into spreading lobules. Column narrow, bended, with 1 stigma and narrow

winged rostellar arms; anther narrowly ovate acute. Pollinium sectile, very long, to 5

mm long, with stipe and viscidium.

3(1) species. Vietnam, Malacca Peninsular, Java, Sumatra, Philippines.

O. echinatum Aver. et Averyanova, 2006, Komarovia, 4: 29, fig. 13.

Described from southern Vietnam (“Thua Thien Hue Prov., Phu Loc Distr.,

Bach Ma national park, S slope of Bach Ma mt., below Hai Vong Dai peak”). Type

(“HLF 1296”) – HN (holotype), LE (isotype).

Stem 20–25 cm tall, with 4–6 uniformly green petiolate leaves, 8–13 cm long,

2.5–4 cm wide. Scape 6–9 cm long, with 2–4 yellowish, broad, acuminate bracts 3–

1.5 cm long; scape, rachis, floral bracts, ovary and sepals glandular hairy. Rachis 7–

9cm long with numerous sessile, odorless flowers, 1.8–2 cm across. Floral bracts

lanceolate, 11–18 mm long. Sepals greenish to olive-pink, sub-similar, ovate, 10–12mm

long; median sepal hooded, forward directed; lateral oblique at the base, spreading.

Petals thin, hyaline, white, oblique-obovate, 9–10 mm long, connate in the apical third.

Lip 9–10 mm long, white. Spur globular, 3–3.5 mm across, inside on each side with

very large gland densely covered with numerous long thick papillae. Base of lip with

erect semi-circular crisped side lobes and 2 semi-circular erect parallel keels. Mesochile


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147

irregularly grooved on sides, narrowing toward epichile. Epichile lobes spreading, flat,

rectangular to obovate, 1.5–3 mm long, 1.5–2 mm wide, irregularly serrate along margin.

Column simple, narrow, 9–10 mm long, bent forward at the middle, with narrowlyovate

stigma at front. Anther and pollinarium 6 mm long. Fig. 40, a-g; 43, h.

Ecology. Coniferous, mixed and broad-leaved evergreen forests on shale and

sandstone, commonly along shady streams. 1000–1100 m. Fl. April – May. Very rare (EN).

Distribution. Vietnam (Thua Thien – Hue).

Note. Species is known up to now only by alone type collection.

Rhomboda Lindl. 1857, Journ. Proc. Linn. Soc. Bot. 1: 181.

Type: Rhomboda longifolia Lindl.

Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome

rooting at nodes, ascending erect leafy floriferous stem and terminal few- to manyflowered

spike. Leaves few, spirally arranged in basal part of stem, petiolate, sheathing

at the base, glabrous, convolute, asymmetric, velvety dark green, brown-green to nearly

black, often with white or pink longitudinal median stripe. Scape leafless, with 2–4

sterile yellowish bracts. Flowers sessile, resupinate or not. Sepals free. Petals oblique,

connivent to median sepal forming hood, lateral sepals spreading. Lip adnate to column

base, broadly saccate at the base. Hypochile with 2 short large peltate glands on

lateral sides, rounded side lobes and 2 prominent lamellate, closely parallel keels,

sometimes with a low keel on flanges. Mesochile channeled, clawed, usually very

short. Epichile entire to 2-lobed. Column clavate, with 2 wings at front, 2 lateral stigmas

and slender rostellum. Anther ovate, acute. Pollinia 2, sectile, with stipe and viscidium.

22(3) species. Bhutan, Nepal, NE India, S China, Myanmar, Thailand, Malaysia,

Indonesia, New Guinea, Solomon Islands, NE Australia.

Key to species

1. Flowers more than 5 mm across; lip 9–12 mm long; sepals dark purple-brown .....................

................................................................................................................................ 1. R. petelotii

– Flowers less than 5 mm across; lip less than 5 mm long; sepals pale brown or greenish

.................................................................................................................................................. 2

2. Epichile entire, broadly-ovate to circular; ovaries erect, densely adpressed to the

rachis ......................................................................................................................... 2. R. tokioi

– Epichile 2-lobed, with spreading acute lobes; ovaries spreading, not adpressed to the

rachis ................................................................................................................ 3. R. lanceolata

1. R. petelotii (Gagnep.) Ormerod, 1998, Austral. Orchid Rev. 63: 11; Aver.

et Averyanova, 2003, Updated Checklist Orch. Viet.: 11. – Haemaria petelotii Gagnep.

1933, Bull. Soc. Bot. France, 80: 349. – Anoectochilus petelotii (Gagnep.) Seidenf.

1975, Contr. Revis. Orch. Fl. Cambodia. Laos Vietnam: 7. – Ludisia discolor auct.

non (Ker Gawl.) A. Rich.: Seidenf. 1992, Orch. Indochina: 29, p. p.

Described from northern Vietnam (“Tonkin: Cha-pa, 1.500–1.600 m.”). Type

(“Petelot 5166”) – P (holotype), C (isotype).


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Terrestrial and lithophytic herb 20–35 cm tall with 3–5 petiolate leaves in basal

rosette. Leaves ovate, 4–7 cm long, acute velvety dark green, purple-brown to nearly

black, with 1 (rarely 3) white to light pink longitudinal stripes. Scape 7–12 cm tall,

shortly pubescent, with 2–3 cuneate, acuminate, yellowish bracts. Rachis 6–15 cm

long, with many not resupinate flowers. Ovary glabrous, 10–15 mm long. Floral bracts

sparsely pubescent at the base, ciliate along margin, narrowly-ovate, acuminate, as

long as, or slightly shorter than ovary. Sepals purple-brown, free, sparsely pubescent

at the base; median sepal narrowly ovate, acute, 7–9 mm long, with recurved apex;

lateral sepals oblique-ovate, narrowing to the apex, 8–10 mm long. Petals brown,

narrowly-ovate, oblique, falcate, acuminate, as long as median sepal. Lip 9–10 mm

long. Hypochile purple to purple-brown, saccate, 5–6 mm long, 4–5 mm wide, with 2

roundish, erect side lobes, 2 large stalked hemi-circular gland at the base, 1 short tall

triangular lamellate keel along midrib and lamellate, finely serrulate lateral longitudinal

flanges outside. Epichile joined to hypochile with a short claw, sometimes twisted on

180 о . Epichile lobes circular, spreading, irregular wavy, 5–6 mm across. Column conical,

4–5 mm tall, with 2 large broad, triangular wavy wings at front, acute bifid rostellum

and 2 lateral stigmas touching at front. Anther acuminate at the apex. Fruits ellipsoid

capsule, 12–15 mm long, 3–5.5 mm wide. Fig. 40, h-l; 43, i.

Ecology. Humid forests on rich soils, mainly on rocky limestone. 700–2000 m.

Fl. July – August (VU).

Distribution. Vietnam (Bac Kan, Cao Bang, Ha Giang, Hoa Binh, Kon Tum,

Lai Chau, Lam Dong, Lao Cai, Quang Binh, Son La, Thanh Hoa, Thua Thien – Hue).

Studied specimens. Bac Kan, Ba Be national park, HLF 514 (HN, LE), HLF 552

(HN, LE); Bac Kan, Cho Don, HLF 761 (HN, LE), HLF 804 (HN), HAL 4790 (HN, LE); Bac Kan,

Na Ri, HAL 4924 (HN); Cao Bang, Bao Lac, CBL 312 (HN, LE); Cao Bang, Tra Linh, CBL 983

(HN, LE); Cao Bang, Trung Khanh, HAL 5561 (HN, LE); Ha Giang, Bac Me, HAL 6577 (HN,

LE); Ha Giang, Meo Vac, NTH 3442 (HN, LE); Ha Giang, Quan Ba, NTH 3603 (HN, LE); Ha

Giang, Yen Minh, NTH 3459 (HN, LE), NTH 3459 (HN, LE); Hoa Binh, Da Bac, HAL 467/a (HN,

LE); Hoa Binh, Mai Chau, HAL 641 (HN, LE), HAL 690 (HN, LE), DKH 7716 (H, LE, MO), DKH

7921 (HN, LE, MO); Kon Tum, Dak Gley, VH 1850 (HN, LE); Kon Tum, Kon Plong, VH 5439

(HN); Lai Chau, Sin Ho, HAL 9962 (HN), HAL 10050 (HN); Lai Chau, Tam Duong, HAL 10254

(HN); Lam Dong, Lac Duong, Hon Giao ridge, VH 4132 (HN, LE); Lao Cai, Van Ban, HAL 2013

(HN, LE), HAL 2316 (HN), HAL 2656 (HN, LE); Quang Binh, Bo Trach, HAL 6059 (HN, LE),

HAL 6141 (HN, LE photo); Son La, Moc Chau, DKH 7322 (HN, LE, MO), DKH 7357 (HN, LE,

MO), DKH 7452a (HN, LE, MO), DKH 7453 (HN, LE, MO); Son La, Moc Chau, HAL 9304

(HN); Son La, Thuan Chau, HAL 9632 (HN); Son La, Yen Chau, DKH 7204 (HN, LE, MO), HAL

9435 (HN); Thanh Hoa, Ba Thuoc, HAL 3495 (HN, LE); Thua Thien – Hue, A Luoi, HAL 7682

(HN), HAL 7683 (HN, LE); Thua Thien – Hue, Bach Ma national park, HLF 943 (HN, LE), HLF

1002 (HN, LE).

Note. Taxonomically very isolated species, which differs in touching stigma

lobes, serrulate lamellate flanges of mesochile, large alone median keel on the lip disc

and usually twisted claw. The plant is not rare in limestone areas of northern Vietnam

and may be certainly found in allied limestone regions of Laos and China.


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149

2. R. tokioi (Fukuyama) Ormerod, 1998, Austral. Orchid Rev. 63, 4: 11. –

Hetaeria tokioi Fukuyama, 1934, Bot. Mag. (Tokyo) 48: 434. – H. pauciflora auct.

non Ridl.: Seidenf. 1992, Orch. Indochina: 39; Aver. 1994, Identif. Guide Vietnam.

Orch.: 33. – Rhomboda pauciflora auct. non (Ridl.) Ormerod: Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 54.

Described from Taiwan (“Taiwan – Taikohu Praef., near Urai”). Type (09–

1932, Suzuki-Tokio 4128) – TI?

Terrestrial herb 8–17 cm tall. Leaves shortly petiolate, 2–4, in basal rosette,

narrowly-ovate to ovate, acute, 2–6 cm long, velvety green with indistinct median

whitish stripe. Scape shortly tomentose, 4–7 cm tall, with 2–3 cuneate, acuminate

bracts. Rachis 2.5–7 cm long, with numerous lax hardly opening flowers. Floral bracts

narrowly-ovate, acute, sparsely hairy and ciliate along margin, 5–10 mm long. Ovary

erect, adpressed to the rachis, glabrous, longer than floral bracts. Sepals greenishbrown,

similar, ovate, 3.5–4.5 mm long, obtuse, glabrous. Petals white, oblanceolate,

blunt at the apex, as long as sepals. Lip white, ovate, concave, as long as sepals.

Hypochile with 2 stalked, warty glands at the base and 2 short semi-circular keels

inside. Epichile ovate to sub-circular, 0.8 mm across. Column conical, 2.5–3 mm tall,

with 2 parallel keels at front and narrow bifid acute rostellum. Anther narrowly ovate,

acuminate. Fig. 40, m, n; 44, a.

Ecology. Humid broad-leaved lowland forests on silicate soils. 400–450 m.

Fl. May – June. Rare (DD).

Distribution. Vietnam (Ha Tinh), Japan, SE China, Hainan, Taiwan.

Studied specimens. Ha Tinh, Huong Son, HAL 5216 (LE); Vietnam, sine loc.,

1985, Averyanov et al. LX-VN s.n. (LE).

Note. Available specimens from Vietnam differ in glabrous ovary and sepals

and may be present separate variety.

3. R. lanceolata (Lindl.) Ormerod, 1995, Orchadian, 11: 329 – Dossinia lanceolata

Lindl. 1857, Journ. Proc. Linn. Soc. Bot. 1: 186. – Anoectochilus pomrangianus

Seidenf. 1978, Dansk Bot. Ark. 32, 2: 41; Aver. et Averyanova, 2003, Updated

Checklist Orch. Viet.: 11; N.T. Tich, 2001, Orch. Viet Nam: 178.

Described from NE India (“India, Khasia, Pomrang”). Type (Hooker & Thomson

s.n.) – K.

Herb 10–25 cm tall with glabrous stem. Leaves 3–4, distant on 1.5 –2 cm,

petiolate, lanceolate, acute, 3–6 cm long. Scape 7–8 cm tall, with 2–3 lanceolate,

acute sterile bracts and 10–14 lax flowers. Rachis 5–6 cm long. Floral bracts lanceolate,

ciliate along margin, 3–9 mm long. Ovary twisted, 6–9 mm long. Flowers sub-globose,

about 3 mm across. Sepals similar, pale brown, ovate, acute, 3.5–4.5 mm long. Petals

white, oblique-ovate, obtuse, about 4 mm long. Lip white with pale yellowish base,

longer than sepals, saccate at the base, 4–4.5 mm long, 3 mm wide; sac with 2 lamellate

keels along midrib and large callus on each side; apex decurved, 2-lobe, with central

apiculus; epichile lobes oblique-ovate. Column 2–3 mm tall. Pollinia clavate, 2 mm

long. Fig. 41, a.


150

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 40. Orchipedum echinatum: a – flowering plant, b – flower, c – flattened tepals, d – lip, side

view and lip longitudinal section, e – spur cross section and spur glands, f – column, side and

frontal views, g – pollinarium; Rhomboda petelotii: h – fruiting plant, i – flattened tepals and lip,

j – lip, without side lobe, k – column, side view and longitudinal section of lip base, l – column,

frontal view; R. tokioi: m – flattened flower, without column, n – column, side and frontal views.


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151

Ecology. Mountain forests. 1500 m. Fl.? (DD).

Distribution. Vietnam (Lam Dong), NE India, Malaysia, Sumatra, Sulawesi,

Philippines.

Note. The species is reported according to data of Nguyen Thien Tich (l.c.),

who found it in Lam Dong province, Langbian mountain. Voucher herbarium specimens

were not preserved. Occurrence of species in Vietnam needs confirmation.

Vrydagzynea Blume, 1858, Coll. Orch. Arch. Ind. Jap.: 71. – Vridagzenia

Benth. 1881, Journ. Linn. Soc. London (Bot.) 18: 344, orth. var. rej.

Lectotype: V. albida (Blume) Blume (Etaeria albida Blume)

Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome

rooting at nodes, ascending erect leafy floriferous stem and terminal few flowered,

rather dense spike. Leaves few, spirally arranged, petiolate, sheathing at the base,

more or less distant or crowded at the stem apex, glabrous, convolute, asymmetric,

green to purple, sometimes with median whitish stripe, acute. Scape short, leafless,

with 1–2 sterile bracts. Flowers small, sessile, resupinate, not widely opening. Sepals

free, very fleshy, succulent. Petals connivent to median sepal forming hood. Lip simple,

ovate, fleshy, conical, with spur. Spur saccate to cylindric, with 2 glands at the base.

Column short, erect, with short bifid rostellum, wingless or with 2 insignificant wings

at front. Anther ovate, obtuse. Pollinia 2, sectile, with caudicle and viscidium. Stigma

with 2 separated lobes.

35(1) species. Tropical and subtropical Asia, northeastern Australia to islands

of W Pacific.

V. formosana Hayata, 1916, Icon. Pl. Formos. 6: 88. – V. albida auct. non

(Blume) Blume, 1858: Seidenf. 1992, Opera Bot. 114: 30; Aver. 1994, Ident. Guide

Vietnam. Orch.: 34; Aver. et Averyanova, 2003, Updated Checkl. Orch. Vietnam: 61.

Described from Taiwan (“Inter Urai et Agioku …“). Type (“leg. D. Hayata,

Mai. 1916”) – TI?

Herb 4–20 cm tall. Leaves 4–6, petiolate, ovate, 2–4 cm long, uniform green,

acute. Scape sparsely pubescent, ebracteate or with 1–3 acuminate bracts, 2–5 mm

long. Rachis densely tomentose, 1–3 cm long, elongate in fruits up to 12 cm, with 3–20

close flowers. Floral bracts narrowly-ovate, acuminate, 6–10 mm long. Ovary 5–8

mm long. Flowers hardly opening, 3–4 mm long, white, later yellowish, base of sepals

and spur green to olive-green, later dull orange to red. Sepals sub-similar, narrowlyovate,

very fleshy, 3.5–5 mm long, 1.5–2 mm wide, blunt. Petals lanceolate, of the

same length, connivent to dorsal sepal. Lip 3–3.5 mm long, spurred, fleshy and finely

papillose along midvein, attenuate to short fleshy blunt tip, lateral sides thin incurved;

lip blade being flattened broadly ovate to almost orbicular. Spur broadly-conic, 3.5–4

mm long, with 2 stalked glands. Fruit erect elliptic capsule, 8–10 mm long, 3 mm wide.

Fig. 41, b; 44, b.

Ecology. Humid forests on any kind of soils, commonly along wet, shady

stream and river valleys. 300–1100 m. Fl. January – April. Not rare (LR).


152

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Distribution. Vietnam (Ha Tinh, Hoa Binh, Lao Cai, Nghe An, Ninh Binh,

Quang Binh, Quang Tri, Thua Thien – Hue, Tuyen Quang), SE China, Taiwan, Laos.

Studied specimens. Ha Nam Ninh, Cuc Phuong, sine date, Bong 146 (HN), LX-VN

1767 (LE); Ha Tinh, Huong Son, HAL 5215a (HN, LE); Hoa Binh, Da Bac, HAL 440 (HN, LE);

Lao Cai, Van Ban, DKH 6678a (HN, LE, MO), HAL 2748 (HN, LE); Nghe An, Tuong Duong,

HLF 6738 (HN), HLF 6930 (HN); Quang Binh, Bo Trach, HAL 6058 (HN, LE), HAL 6148 (HN,

LE), HAL 6229 (HN, LE); Quang Binh, Quang Ninh, HAL 6316 (HN, LE) d-EXSICCATES OF

VIETNAMESE FLORA 0005/HAL6316; Quang Tri, Da Krong, HLF 6192 (HN); Thua Thien –

Hue, Nam Dong, HAL 11129 (HN), HAL 6896a (HN, LE); Tuyen Quang, Na Hang, HAL 221

(HN, LE).

Note. Species closely related to V. albida (Blume) Blume, from which it

differs in hairy rachis, floral bracts and ovary, as well as almost rounded to emarginate

lip (when flattened) with very thin incurved lateral margins. Fairly common species in

northern Vietnam, particularly in wet forests on rocky limestone. Easily overlooked in

botanical studies and collecting due to dwarf habit and unattractive appearance of

flowers. All seen collections from Vietnam belong to this species, which replaces

V. albida (distributed in Myanmar, Thailand, Peninsular Malaysia, Indonesia, Philippines

and New Guinea) in Eastern Indochina, mainland SE China and in Taiwan.

Zeuxine Lindl. 1826, Collect. Bot. App. N 18 (sub “Zeuxina” – orth. var.

rej.); id. 1840, Gen. Sp. Orch. Pl.: 485, nom. cons.

Type: Z. strateumatica (L.) Schleichter (Pterygodium sulcatum Roxb.;

Z. sulcata (Roxb.) Lindl.).

Terrestrial and lithophytic sympodial herbs with creeping succulent rhizome

rooting at nodes, ascending erect leafy floriferous stem and terminal few- to many

flowered spike. Leaves few, spirally arranged, petiolate, sheathing at the base, glabrous,

convolute, asymmetric, uniformly green or with median whitish stripe, acute, rarely

conduplicate, grass-like. Scape leafless, with 2–4 sterile bracts. Flowers small, sessile,

resupinate. Sepals free. Petals connivent to median sepal forming hood, lateral sepals

spreading. Lip spurless. Hypochile concave to saccate, with various papillae on lateral

sides. Epichile reflexed, circular-obovate or 2-lobed, connected to hypochile by a short

claw. Column short, erect, with 2 prominent rostellar arms and 2 wings at front. Anther

narrowly triangular-ovate, acute. Pollinia 2, sectile, with caudicle and viscidium. Stigma

with 2, small, convex, separated lobes.

70(7) species. Africa, tropical and subtropical Asia, Indonesia, Philippines,

Australia, New Guinea, Pacific Islands to Samoa.

Key to species

1. Leaves linear, conduplicate, grass-like ................................................... 1. Z. strateumatica

– Leaves lanceolate to almost circular, plicate .................................................................... 2

2. Epichile entire, circular or obcordate ....................................................... 2. Z. goodyeroides

– Epichile distinctly 2-lobed ................................................................................................ 3

3. Epichile with divergent lobes 6–8 mm long; sepals 7–8 mm long ................. 3. Z. grandis

– Epichile with spreading lobes less than 3 mm long; sepals less than 6 mm long ........ 4


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4. Column with large, protruding quadrate or triangular flat vertical wings on front;

petals very oblique, suddenly narrowing toward the obtuse apex ................ 4. Z. nervosa

– Column with insignificant, hardly visible wings on front; petals not much oblique

narrowing to the apex ......................................................................................................... 5

5. Lip white, 3–4 mm long; epichile lobes rectangular, about twice as long as wide ...........

.............................................................................................................................. 5. Z. parvifolia

– Lip yellow, 2–3 mm long; epichile lobes squarish to obovate, less than twice as long

as wide ........................................................................................................................ 6. Z. flava

1. Z. strateumatica (L.) Schleichter, 1911, Bot. Jahrb. 45: 394; Seidenf. 1992,

Opera Bot. 114: 37; Aver. 1994, Identif. Guide Vietnam. Orch.: 38; Aver. et Averyanova,

2003, Updated Checklist Orch. Viet.: 61. – Orchis strateumаtica L. 1753, Sp. Pl.:

943. – Z. bonii Gagnep. 1931, Bull. Mus. Natl. Hist. Nat. 2. ser. 3: 326. – Z. evrardii

Gagnep. 1931, l.c.: 326.

Described from Sri Lanka (“in Zeylonа”). Type (“Hermann s.n.”) – BM.

Stem erect, slender, to 20 cm tall, leafy. Leaves erect, conduplicate, linear,

grass-like, acute, to 6 cm long. Inflorescence many-flowered cylindric dense spike 1–

4 cm long. Ovary glabrous, 5–7 mm long. Floral bracts triangular-ovate, acuminate,

reddish-yellow, 4–10 mm long. Sepals white, sub similar, ovate, obtuse, 3–4 mm long,

glabrous. Petals of the same length, white, oblique narrowly-obovate, obtuse. Lip shortly

saccate, with 2 glands, 3–3.5 mm long; claw short, fleshy, papillose and grooved.

Epichile small, transversely oblong-reniform, papillose, blunt, yellow-green. Column

short, 1.5–2 mm tall. Fig. 41, c-f.

Ecology. Wet secondary grasslands, open wet places along stream and river

valleys. 50–1500 m. Fl. January – February. Very rare (DD).

Distribution. Vietnam (Cao Bang, Ha Noi, Ha Tinh, Lam Dong, Nam Dinh,

Nghe An, Ninh Binh, Thanh Hoa). Tropical and subtropical Asia, Japan, Taiwan,

Philippines, Indonesia to New Guinea.

Studied specimens. Cao Bang; Ha Noi, Kien-Khe, mont Den; Ha Son Binh, Rocher

de Notre Dame, riviere Noire; Lam Dong, Dalat, Camly; Lang-bian; Nghe Tinh; Thanh Hoa;

Tonkin, delta du Fleuve Rouge; Tonkin, Mong-hаo, Fleuve Rouge.

Note. Very variable and widespread species escaped and naturalized in Florida

and Cuba. Recently it, probably, become much rarer as did not been collected in

Vietnam during last 50 years.

2. Z. goodyeroides Lindl. 1840, Gen. Sp. Orch. Pl.: 486.– Z. bidupensis

Aver. 2006, Rheedea, 16, 1: 12, fig. 7g, h & 9.

Described from NE India (“India, Assam ...”). Type (“Griffith 7”) – K.

Terrestrial herb. Stem 15–20 cm tall with 4–6 leaves. Petiole and sheath 5–10

mm long; leaf blade ovate to narrowly-ovate, acute, 2–4 cm long, dark velvety green

to green-brown, sometimes with median whitish stripe. Scape and rachis sparsely

pubescent, rachis lax flowered, commonly with 3–10 flowers. Floral bracts dull reddishbrown,

triangular-ovate, acuminate, glabrous, erose-ciliate along the margin, 4–7 mm

long, 2–3 mm wide. Ovary dark olive-green, cylindric, glabrous, 8–10 mm long. Flowers


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

odorless. Sepals olive-brown, glabrous, sub-similar, narrowly-ovate, acute, 5.5–6 mm

long; dorsal sepal wider. Petals white, oblique, falcate, rising from narrow base to

oblique broad obtuse apical part, 5–5.5 mm long. Lip 4–4.5 mm long, 2.5–3 mm wide.

Hypochile yellowish, ovate, concave, 2.5–3 mm long, 2.5–3 mm wide, with 2 swollen

nerves and 2 hooked or indistinctly T-shaped glands inside; margins incurved, fleshy,

coming apically into 2 papillose swellings. Epichile white, entire, circular or obcordate,

1–1.2 mm long, 1.2–1.4 mm wide, with finely crenulate margin. Column 2.5–3 mm

tall, with 2 short triangular acute wings at front and 2 long narrow falcate apical

rostellum arms. Anther narrowly-ovate, acuminate, 2.2–2.5 mm long. Pollinia massulae

brown, triangular, 0.1 mm across. Fig. 41, g-k; 44, c.

Ecology. Wet broad-leaved and coniferous closed forests on silicate soils.

1500–1900 m. Fl. November – January. Rare (VU).

Distribution. Vietnam (Lai Chau, Lam Dong), Bhutan, NE India, Myanmar,

S China.

Studied specimens. Lai Chau, Phong Tho, HAL 10670 (HN, LE), HAL 10672 (HN,

LE); Lam Dong, Lac Duong, Gia Rich mt., HLF 5319 (HN, LE).

3. Z. grandis Seidenf. 1978, Dansk Bot. Ark. 32: 90. – Z. affinis auct. non

(Lindl.) Benth. ex Hook. f.: Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 61.

Described from Thailand (“Omkoi Road at km 13”). Type (“GT 7730”) – C.

Terrestrial and lithophytic herb. Stem 8–25 cm tall with 2–6 leaves, usually

withered at the time of flowering. Leaves petiolate; narrowly-ovate, 1.5–4 cm long,

uniform dull green, acute. Scape and rachis densely pubescent, rachis to 6 cm long,

with 1–10 lax flowers. Floral bracts triangular-ovate, acuminate, pubescent, 6–12 mm

long. Ovary pubescent, 5–9 mm long. Sepals olive-green to light pinkish, pubescent

outside, sub-similar, ovate, 7–8 mm long, concave, obtuse. Petals of the same length

or longer, white, oblique narrowly-obovate to oblong-spathulate, roundish at the apex.

Lip white, with light yellow-brownish central spot on epichile, 10–13 mm long. Hypochile

small, saccate, with 1 small gland on each lateral side. Epichile 2–lobed, lobes flat, at

an acute angle to the axis, convergent, oblique-obovate, each 6–8 mm long, 3.5–

4.5 mm wide. Column short, without frontal wings. Fruit erect elliptic capsule, 8–

12 mm long, 2–3 mm wide. Fig. 41, l, m; 44, d.

Ecology. Wet primary and secondary closed forests, often with bamboo, on shale,

usually along rocky stream slopes. 1000–1200 m. Fl. February – March. Rare (EN).

Distribution. Vietnam (Quang Tri), Thailand, Hainan.

Studied specimens. Quang Tri, Huong Hoa, Sa Mui pass, HLF 5755a (HN, LE),

HLF 6278 (HN, LE); d-EXSICCATES OF VIETNAMESE FLORA 0069/HLF6278.

4. Z. nervosa (Wall. ex Lindl.) Benth. ex Clarke, 1889, Journ. Linn. Soc.

London (Bot.) 25: 73; Seidenf. 1992, Opera Bot. 114: 37; Aver. 1994, Identif. Guide

Vietnam. Orch.: 38; Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 61. –

Monochilus nervosum Wall. ex Lindl. 1840, Gen. Sp. Orch. Pl.: 487.

Described from E India (“India orientali”). Type (“Wallich 7381 А”) – K?


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155

Terrestrial herb. Stem 10–25 cm tall with 3–5 leaves. Leaves petiolate; petiole

and sheath 0.5–1.5 cm long; leaf blade ovate to narrowly-ovate, 2–6 cm long, pure dull

green, often with wide median whitish stripe, acute. Scape and rachis sparsely pubescent,

rachis 1–6 cm long, with 3–14 laxy flowers. Floral bracts triangular-ovate, acuminate,

5–10 mm long. Ovary sparsely hairy, 5–10 mm long. Sepals dull greenish to greenishbrown,

sub-similar, ovate, 3.5–5.5 mm long, concave, obtuse. Petals of the same length,

white, oblique, broadly lunate, attenuate at the apex. Lip white, with 2 dull green to

brown spots on epichile, 4.5–6 mm long. Hypochile saccate, with (1)2–3 close parallel

fat calluses on each lateral side. Epichile 2-lobed, lobes spreading, rectangular to obovate.

Column short, with 2 large rectangular to triangular acute wings at front. Fruit erect

elliptic capsule, 9–12 mm long, 3.5–4.5 mm wide. Fig. 41, n, o; 44, e.

Ecology. Wet primary and secondary closed forests, often with bamboo, on

any soils. 200–1500 m. Fl. January – April. Not rare (LR).

Distribution. Vietnam (probably all provinces), Bhutan, Nepal, India, S China,

Thailand, Ryukyu Islands, Taiwan, Philippines.

Studied specimens. Cuc Phuong national park, NTH 3890 (HN, LE); Ha Giang,

Hoang Su Phi, HAL 6619 (HN, LE); Ha Giang, Quan Ba, DKH 5008 (HN, LE, MO); Ha Nam

Ninh, Cuc Phuong national park, LX-VN 1810 (HN, LE); Ha Tinh, Huong Son, HAL 5215a

(HN, LE); Hoa Binh, Da Bac, HAL 393 (HN, LE); Lao Cai, Van Ban, HAL 2001 (HN, LE); Nghe

An, Quy Chau, HLF 3095 (HN, LE); Nghe An, Tuong Duong, HLF 6602 (HN, LE); Quang

Binh, Minh Hoa, HAL 5868 (HN, LE); Son La, Moc Chau, DKH 7357 (HN, LE, MO); Son La,

Yen Chau, DKH 7225 (HN, LE, MO); Tuyen Quang, Na Hang, HAL 181 (HN, LE).

Note. Widespread and common species in Vietnam, which easily survives in

wet secondary forests, particularly with bamboo.

5. Z. parvifolia (Ridl.) Seidenf. 1978, Dansk Bot. Ark. 32, 2: 82; Seidenf.

1992, Opera Bot. 114: 37; Aver. 1994, Identif. Guide Vietnam. Orch.: 37; Aver. et

Averyanova, 2003, Updated Checklist Orch. Viet.: 61. – Hetaeria parvifolia Ridl.

1903, Journ. Straits Branch Roy. As. Soc. 39: 87. – Z. tonkinensis Gagnep. 1931,

Bull. Mus. Natl. Hist. Nat. 2. s. 3, 7: 328.

Described from Malacca Peninsula (“Penins. Mal.” – sec. Ind. Kew.). Type

(“Ridley s.n.”) – K.

Terrestrial herb. Stem 10–30 cm tall with 3–5 leaves. Leaves shortly petiolate,

ovate to narrowly-ovate, 1.5–4.5 cm long, uniform dull green, or sometimes with median

whitish stripe, acute. Scape and rachis sparsely pubescent, rachis 1–8 cm long, fewto

many flowered. Floral bracts triangular-ovate, acuminate, sparsely pubescent to

glabrous, 5–10 mm long. Ovary sparsely pubescent or glabrous, 4–8 mm long. Sepals

light greenish-brown, sub-similar, ovate to narrowly-ovate, 3–4.5 mm long, concave,

obtuse. Petals of the same length, white, oblique-oblong, roundish at the apex. Lip 3–

4 mm long, white, with yellowish to yellowish-brown spot on epichile. Hypochile saccate,

with 1 small gland on each lateral side. Epichile 2-lobed, lobes spreading, rectangular,

about twice as long as wide. Column short, with 2 narrow insignificant wings at front.

Fig. 41, p; 44, f.


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 41. Rhomboda lanceolata: a – flower and floral bract; Vrydagzynea formosana: b – flattened

flower and longitudinal section of spur; Zeuxine strateumatica: c – flowering plant, d – flower, e –

flattened sepals and petals, f – column and lip, side view; Z. goodyeroides: g – flowering plant,

h – flattened tepals and lip, i – column, frontal fiew, j – anther, ventral view, k – ovary and floral bract;

Z. grandis: l – flattened tepals and lip, m – column, frontal view; Z. nervosa: n – flattened tepals

and lip, o – column, side and frontal views; Z. parvifolia: p – column, flattened tepals and lip.


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157

Ecology. Wet primary and secondary closed forests, often with bamboo, on

any soils. 300–1800 m. Fl. December – April. Not rare (LR).

Distribution. Vietnam (probably all provinces), Myanmar, Thailand, Laos,

Cambodia, Malacca Peninsula, Ryukyu Islands, Taiwan, Philippines.

Studied specimens. Bavi mt., Tonkin, Balansa 2002 (P); Dak Nong, Dak Glong,

HLF 5503a (HN, LE); Gia Lai – Kontum, Chu Pa, LX-VN 2531 (HN, LE); Ha Giang, Hoang Su

Phi, HAL 6620 (HN, LE); Hoa Binh prov., Da Bac, HAL 556 (HN, LE); Kon Tum, Dak Gley, VH

2281 (HN, LE); Lam Dong, Dalat, VH 2523 (HN, LE); Lam Dong, Lac Duong, Bi Dup mt, VH

3236 (HN, LE); Lang Son, Bac Son, HAL 6790 (HN, LE); Lao Cai, Van Ban, HAL 2265 (HN, LE),

DKH 6740 (HN, LE, MO); Thanh Hoa, Ba Thuoc, HAL 3310a (HN, LE); Thua Thien – Hue,

Nam Dong, HAL 6896b (HN, LE).

Note. Widespread and common species in Vietnam, which easily survives in

wet secondary forests, particularly with bamboo.

6. Z. flava (Wall. ex Lindl.) Trimen, 1885, Syst. Cat. Fl. Pl. Ceylon: 90. –

Monochilus flavus Wall. ex Lindl. 1840, Gen. Sp. Orch. Pl.: 487.

Described from Nepal. Type (“Wallich, Wall. Cat. 7380A”) – K.

Terrestrial and lithophytic herb. Stem 15–45 cm tall with 5–8 leaves. Leaves

petiolate; ovate to narrowly-ovate, 4–8 cm long, uniform light green, acute. Scape and

rachis pubescent, rachis to 22 cm long, many flowered. Floral bracts triangular-ovate,

acuminate, sparsely pubescent, 5–11 mm long. Ovary sparsely pubescent to subglabrous,

6–10 mm long. Flowers odorless. Sepals dull greenish-brown, sub-similar,

ovate to narrowly-ovate, 3.5–4 mm long, concave, obtuse, sparsely pubescent to glabrous,

lateral sepals shorter. Petals of the same length, light greenish, oblique narrowly-obovate

to oblong-lanceolate, obtuse. Lip yellow, 2–3 mm long. Hypochile saccate, with 1

small gland on each lateral side. Epichile 2-lobed, lobes spreading, oblong-obovate,

0.8–1.2 mm long. Column short, with 2 narrow, fat, insignificant wings at front. Fruit

erect elliptic capsule, 6–9 mm long, 2–3 mm wide. Fig. 42, a, b; 44, g.

Ecology. Wet primary and secondary closed forests, often with bamboo, on

any soils. 500–1500 m. Fl. February – April. Rare (VU).

Distribution. Vietnam (Ninh Thuan, Quang Tri, Son La), Bhutan, Nepal, N

India, Myanmar, Thailand.

Studied specimens. Ninh Thuan, Ninh Hai, Nui Chua mt, HLF 4445 (HN, LE),

HLF 4538 (HN, LE); Quang Tri, Huong Hoa, Sa Mui pass, HLF 5939 (HN, LE); Son La, Moc

Chau, DKH 7515 (HN, LE, MO).

Zeuxinella Aver. in Aver. et Averyanova, 2003, Updated Checklist Orch. Viet.: 96.

Type: Z. vietnamica (Aver.) Aver. (Zeuxine vietnamica Aver.)

Terrestrial creeping sympodial herb with succulent rhizome rooting at nodes,

short ascending leafy stem and erect thin terminal leafless inflorescence 10–18 cm

tall. Leaves 3–5, petiolate. Petiole and sheath 0.5–1 cm long. Leaf blade velvety

carbon-black above, pale dull purple below, broadly ovate to orbicular, 1.5–3 cm long,

shortly apiculate. Scape hairy, with 3–4 acuminate bracts; rachis pubescent, 1–4 cm


158

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

long, with 3–8 laxy sessile flowers. Floral bracts pubescent, cuneate, acuminate, 4–6

by 1–1.5 mm. Ovary densely glandular hairy, 5–7 mm long. Flowers hardly opening,

odorless, resupinate, 3–5.5 mm long. Sepals dull greenish to greenish-brown, subsimilar,

narrowly-ovate to ovate, 3–5 mm long, concave, obtuse. Petals white, narrowly

ovate, incurved, 3–5 mm long, connivent to dorsal sepal forming hood. Lip yellow,

spurred, fleshy, 3-lobed, 3 mm long. Hypochile boat-shaped; sidelobes narrowly ovate,

rising and incurved. Epichile reflexed, 3-lobulate, with rounded-triangular subequal

lobules, with 4 cushions on the disk. Spur straight, sac-like, 2.5–3 by 2 mm, with 1

cylindric gland on each lateral side. Column small, with 2 insignificant frontal wings, 2

narrow rostellar arms and 2 stigmas. Fig. 42, c-h; 44, h.

Monotypic genus endemic to northern Vietnam.

Z. vietnamica (Aver.) Aver. 2003, in Aver. et Averyanova, Updated Checkl.

Orch. Vietnam: 61, 96, fig. 11. – Zeuxine vietnamica Aver. 1988, Bot. Journ. (Leningrad)

73, 3: 424; Seidenf. 1992, Opera Bot. 114: 38.

Described from northern Vietnam (“Prov. Ha Nam Ninh. Cuc Phuong, May

bac”). Type (“2 May 1985 LX-VN 1809/52”) – HN (holotype), LE (isotype).

Ecology. Primary and secondary broad-leaved forests, often with bamboo,

on rocky limestone. 50–400 m. Fl. March – May. Very rare (EN).

Distribution. Vietnam (Bac Kan, Ninh Binh, Quang Ninh).

Studied specimens. Bac Kan, Ba Be national park, HLF 638 (HN, LE); Ha Nam

Ninh, Cuc Phuong, NTH 3891 (HN, LE); Quang Ninh, Ha Long Bay, NTH 5649 (HN, LE).

Notes. Very rare relictual element of lowland indigenous limestone floras of

northern Vietnam. The only representative of monotypic strictly endemic genus that is

one of the best examples of strict endemism at the genus level. Needs special attention

for protection.

Subtrib. 3.2.2. Spiranthinae Lindl.

1840, Gen. Sp. Orch. Pl. : 441, 462 (sub “Spiranthidae”).

Type: Spiranthes L.C. Rich.

28(1) genera and 275(1) species. Tropical, subtropical and temperate regions

of all continents except Africa, with highest diversity in tropical America.

Spiranthes L.C. Rich. 1817, Orch. Europ. Annot.: 20, nom. cons.

Type: S. spiralis (L.) K. Koch (Ophrys spiralis L.), type cons.

Terrestrial herbs with few fleshy fasciculate root tuberoids, rosulate sessile

conduplicate leaves and erect inflorescence bearing terminal many-flowered, secund,

spirally twisted, dense spike. Flowers sessile, resupinate, not widely opening. Sepals

free, sub-similar; median sepal erect, lateral sepals spreading, oblique at the base.

Petals connivent to sepal forming hood. Lip sessile or shortly claved, entire or lobed,

undulate along margin, enveloping column at the base. Column short, erect, with bifid

rostellum and 1 convex, entire stigma at front; clinandrium with prominent margins.

Anther erect, dorsal. Pollinia 2, mealy, with caudicle and viscidium.


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159

Fig. 42. Zeuxine flava: a – flattened tepals and lip, b – column, frontal view; Zeuxinella

vietnamica: c – flowering plant, d – ovary, floral bract and flower, e – flattened tepals and lip,

f – lip, side view, g – gland inside spur, h – column, view from above (with removed operculum)

and frontal view; Spiranthes sinensis: i – flowering plants, j – flower, k – flattened tepals and

lip, l – flattened median sepal and petals, m – flattened lip, n – ovary, column and lip, side view,

o – column, side view, p – column, view from above and frontal view, q – fruit, r – floral bract.


160

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

40(1) species. Tropical, subtropical and temperate areas of the world except

tropical and southern Africa.

1. S. sinensis (Pers.) Ames, 1908, Orch. 2: 53; Seidenf. 1992, Opera Bot.

114: 25; Aver. 1994, Identif. Guide Vietnam. Orch.: 48; Aver. et Averyanova, 2003,

Updated Checklist Orch. Viet.: 56. – Neottia sinensis Pers. 1807, Syn. Pl. 2: 511. –

Aristotelea spiralis Lour. 1790, Fl. Cochinchin.: 522. – N. australis R. Br. 1810,

Prodr.: 319. – N. amoena M. Bieb. 1819, Fl. Taur.-Caucas. 3: 606. – S. australis

(R. Br.) Lindl. 1824, Bot. Reg. 10: tab. 823. – S. amoena (M. Bieb.) Spreng. 1826, Syst.

Veg. 3: 708. – S. spiralis (Lour.) Makino, 1926, Journ. Jap. Bot. 3: 25, nom. illeg.

Described from southern China (“prope Gantonem Sinarum”). Type (“Loureiro

s.n.”)– P.

Herb 8–40 cm tall with 3–10 linear-lanceolate to broadly-lanceolate, glossy

green, acute leaves, 2–12 cm long, 3–8 mm wide. Scape glandular hairy, with 2–4

bracts. Rachis 4–14 cm. Floral bracts lanceolate, acuminate, glandular hairy, 4–7 mm

long. Ovary 2–4 mm long. Flowers pink-purple to white. Sepals and petals sub-similar,

narrowly ovate to lanceolate, 3–4 mm long, 1–1.5 mm wide, obtuse, petals slightly

smaller. Lip simple, oblong, longitudinally concave, 3.5–4.5 mm long, 2–3 mm wide,

plicate along margin, with 2 large spherical glands at the base. Column 2–3 mm tall,

with 2 erect, acute rostellar arms. Fruit ovate, glandular pubescent capsule, 6–7 mm

long, 3 mm broad. Fig. 42, i-r; 44, i.

Ecology. Secondary short-tall grasslands, pastures, road-cuts, old rice field in

agricultural areas, open grassy swamps along stream and river valleys, usually on wet

alluvial soils. 5–2000 m. Fl. February – April. Very common (LR).

Distribution. Vietnam (probably all provinces). Widespread in temperate,

subtropical and tropical Asia to Australia, New Zeeland, Tasmania and south-west

Pacific Islands.

Studied specimens. Annam, sine loc., Eberhardt s.n. (P); Bac Kan, Ngan Son,

CBL 1258 (HN, LE); Cao Bang, Bao Lac, CBL 1373 (HN, LE); Cao Bang, Nguyen Binh, CBL

131 (HN, LE); Cao Bang, Tien Ha, sine coll. (P); Cochimchina s.n., Loureiro (P); Dalat, Boden,

Kloss s.n. (BM); Ha Giang, Bac Me, HAL 6607 (HN, LE); Ha Giang, Dong Van, CBL 1772 (HN,

LE); Ha Giang, Quang Ba, DKH 4832 (HN, LE); Ha Giang, Yen Minh, CBL 1979 (HN); Hanoi,

Bat Bac, Balansa; Hanoi, Kien Khe, Bon; Hoa Binh, Da Bac, HAL 285 (HN); Hoa Binh, Da Bac,

HAL 554 (HN); Ja Lao Mahay, Pierre s.n. (P); Kon Tum, Kon Plong, DKH 4750 (HN, LE);

Kontum, Kon Plong, Mang La, VH 5456 (HN); Kontum, Ngoc Linh mt., VH 10 (HN, LE); Lac

Duong, Langbian, sine coll.; Lang Son, Bac Son, HAL 6791 (HN, LE); Langbian, Andre; Lao

Cai, Van Ban, HAL 2448 (HN); Lao Cai, Van Ban, HAL 2845 (HN, LE); Ninh Binh, Cuc Phuong

national park, 20 Feb. 1971, Quynh 41, CPNP 5120 (HN); Pagode de Bamly, Tonkin, Petelot;

Quang Yen, d’Alleizette; Santay, Balansa; Son La, Moc Chau, DKH 7449 (HN); Thai Nguyen,

Eberhardt; Tonkin delta, Eberhard drawing, 218; Tonkin, sine loc., Bon; Tu Phap, Balansa.

Note. Very common, variable and widespread weedy species, often spreading

into secondary habitats on agricultural lands. Used in traditional oriental medicine.


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161

Fig. 43. Ludisia discolor: a (HLF 6570), b (HAL 6350), c (HAL 1340, HAL 1655 and HAL 5865);

Odontochilus saprophyticus: d (HAL 8194); O. elwesii: e (HAL 1720); O. umbrosus: f (HLF

5611a); O. inabai: g (HAL 9002); Orchipedum echinatum: h (HLF1296, type); Rhomboda

petelotii: i (NTH 3459).


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Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Fig. 44. Rhomboda tokioi: a (LX-VN s.n., 1985); Vrydagzynea formosana: b (HAL 6148); Zeuxine

goodyeroides: c (HAL 10672); Z. grandis: d (HLF 6278); Z. nervosa: e (Averyanov, HAL

s.n., 2004); Z. parvifolia: f (HAL 6620). Z. flava: g (HLF 5939); Zeuxinella vietnamica: h (HLF

638); Spiranthes sinensis: i (HAL 11131a).


Turczaninowia 2008, 11(1) : 5–168

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Gagnepain F., A. Guillaumin. Orchidacees, Apostasiacees. In Lecomte, H. (ed). Flore

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Ban Tre, 2000. – 1020 p.

Seidenfaden G. The Orchids of Indochina // Opera Botanica, 1992. Vol. 114. – 502 p.

Tran Hop. The orchids of Vietnam. – Ho Chi Minh. Nha Xuat Ban Nong Nghiep, 1998. – 703 p.

РЕЗЮМЕ

Статья открывает публикацию серийного издания иллюстрированного критического

таксономического обзора орхидных (Orchidaceae) во флоре Вьетнама. Первая часть монографии

включает предисловие, иллюстрированный словарь терминов, используемых при

описании таксонов и составлении ключей для определения, родовой ключ семейства и

стандартную таксономическую обработку трех подсемейств – Apostasioideae, Cypripedioideae

и Spiranthoideae, включающих 21 род и 85 видов. Для всех признаваемых видов и

разновидностей приводятся законное название, наиболее важные синонимы, цитирование

аутентичного материала, краткое описание, данные по экологии и распространению, а

также список изученных образцов. Все виды и разновидности иллюстрированы черно-белыми

рисунками и цветными фотографиями. Для 4 таксонов в работе приняты новые номенклатурные

комбинации в ранге разновидности. Это – Neuwiedia zollingeri var. annamensis

(Gagnep.) Aver., N. zollingeri var. balansae (Gagnep.) Aver., Paphiopedilum barbigerum var.

aspersum (Aver.) Aver. и P. callosum var. warnerianum (T. Moore) P.J. Cribb ex Aver.


164

Acampe 8, 46

Acanthephippum 7, 41

Acianthinae 6

Acriopsidinae 8

Acriopsis 8, 44

Adenoncos 8, 45

Aerides 8, 48

Aeridinae 8, 33

Aetheria anomala 128

Agrostophyllum 7, 43

Allochilus eberhardtii 120

Amitostigma 6, 39

Anoectochilus 6, 39, 95

- acalcaratus 141

- albolineatus 96, 100, 101, 102, 115

- annamensis 96, 100, 101, 115, 116

- brevilabris 96, 102, 103, 108

- brevistylus 141

- calcareus 96, 99, 101, 115

- chapaensis 103

- daoensis 140

- dawsonianus 132

- elwesii 140

- flavescens 138

- gracilis 134

- griffithii 103

- inabai 146

- koshunensis 99

- lanceolatus 143

- lylei 96, 101, 102, 116

- papillosus 96, 97, 98, 99, 115

- petelotii 147

- pomrangianus 149

- repens 144

- roxburghii 96, 98, 115

- setaceus 95

- setaceus 96

- siamensis 99

- sikkimensis 103

- tonkinensis 141

- tortus144

- tridentatus 103

Anthogonium 7, 41

Aphyllorchis 6, 50

Apostasia 6, 37, 50, 52

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

INDEX OF LATIN NAMES

- nipponica 52, 54, 56, 57

- nuda 52, 54, 57

- odorata 14, 52, 53, 54, 56

- thorelii 53, 56

- wallichii 53, 57

Apostasiaceae 37, 50

Apostasioideae 5, 6, 8, 17, 50, 163

Appendicula 7, 43

Arachnis 8, 46

Arethuseae 7

Arisanorchis takeoi 107

Aristotelea spiralis 160

Arundina 7, 40

Arundinae 7

Ascocentropsis 8, 48

Ascocentrum 8, 48

Biermannia 8, 48

Bletia 7, 41

Bletiinae 7

Brachycorythis 6, 39

Brachypeza 8, 48

Bromheadia 8, 44

Bromheadiinae 8

Bulbophyllinae 8

Bulbophyllum 8, 28, 43

Calanthe 7, 15, 41

Callostylis 7, 44

Cephalantheropsis 7, 41

Ceratostylis 7, 43

Cerochilus rubens 131

Chamaegastrodia poilanei 139

Cheirostylis 6, 32, 39, 104

- bipunctata 105, 112, 125

- chinensis 105, 109, 113, 116

- cochinchinensis 105, 110, 113, 116

- eglandulosa 107

- filipetala 105, 109, 113

- flabellatа 110

- foliosa 104, 106, 108, 116

- grandiflora 114

- inabai 104, 106, 108

- latipetala 105, 112, 123, 137

- marmorata 107

- marmorifolia 105, 107, 113, 116

- montana 104

- phamhoangii 106

- pingbianensis 106

- serpens 104, 105, 108, 116

- spathulata 105, 114, 123, 137

- takeoi 104, 107, 108, 116

- thanmoiensis 105, 112, 113

- yunnanensis 105, 111, 113, 116

Chiloschista 8, 47

Christensonia 8, 47

Chrysobaphus roxburghii 96

Chrysoglossum 8, 44

Cleisocentron 8, 47

Cleisomeria 46

Cleisostoma 46, 47

Cleisostomopsis 46

Cnemidia angulosa 94

Coelogyne 7, 15, 42

- cristata 14

- fimbriata 14

- lentiginosa 14

Coelogyneae 7

Coelogyninae 7

Collabium 8, 44

Collabiinae 8

Corybas 6, 39

Corymborkis 6, 38, 92

- corymbis 92

- veratrifolia 93, 98, 115

Cranichideae 6, 95

Cranichis 95

Cremastra 8, 45

Cryptochilus 7, 44

Cryptopylos 8, 48

Cryptostylidinae 6

Cryptostylis 6, 38

Cymbidieae 8

Cymbidiinae 8

Cymbidium 8, 44

Cypripediaceae 37

Cypripedioideae 5, 6, 8, 19, 58, 163

Cypripedium 58

- appletonianum 80

- barbatum var. warnerianum 82

- boxallii 80

- callosum 80


Turczaninowia 2008, 11(1) : 5–168

- callosum var. sublaeve 82

- concolor 67

- delenatii 61

- gratrixianum 76

- hirsutissimum 69

- insigne 58

- purpuratum 83

- sinicum 83

- villosum 77

Cyrtosia 6, 49

Dactylorhiza incarnata 27

Dendrobiinae 8

Dendrobioideae 12

Dendrobium 8, 29, 43

- cariniferum 14

Dendrochilum 7

Didymoplexiopsis 7, 49

Didymoplexiella 7, 49

Didymoplexis 7, 49

Diglyphosa 8, 44

Diplomeris 6, 39

Diploplora 8, 45

Dipodium 8, 44

Diurideae 6

Doritis 8, 45

- pulcherrima 27

Dossinia lanceolata 149

Eparmatostigma 8, 48

Epidendreae 7

Epidendroideae 6, 23

Epigeneium 8, 43

Epipactis 6, 38

Epipogiinae 7

Epipogium 7, 50

Eria 7, 29, 44

Eriinae 7

Eriodes 7, 41

Erythrodes 6, 38, 117

- blumei 117, 118, 123, 137

- chinensis 118

- herpysmoides 117

- hirsuta 117, 123, 137

- latifolia 117

- seshagiriana 117

Erytrorchis 6, 49

Esmeralda 8, 45

Etaeria 128

- albida 151

- oblongifolia 128, 129

Evrardiа 138

- poilanei 139

Evrardianthe 138

- poilanei 139

Evrardianа 138

- poilanei 139

Eulophia 8, 45

Eulophiinae 8

Flickingeria 8, 42

Galeola 6, 49

Galeolinae 6

Gastrochilus 8, 48

Gastrodia 7, 49

Gastrodieae 7

Gastrodiinae 7

Geodorum 8, 45

Georchis biflora 122

- foliosa 124

Glomerinae 7

Goodyera 6, 38, 95, 118

- affinis 131

- arisanensis 119, 126, 130

- biflora 119, 122, 123, 137

- discolor 132

- foliosa 119, 124, 127, 130, 138

- fumata 119, 120, 123, 137

- hirsuta 117

- hispida 119, 121, 123, 137

- procera 119, 120, 123, 137

- repens 118

- rhombodoides 119, 126, 130

- schlechtendaliana 119, 125, 130, 138

- viridiflora 119, 123, 124, 137

Goodyerinae 6, 32, 95

Grossourdya 8, 49

Habenaria 6, 39, 40

- rhodocheila 39

Habenariinae 6

Haemaria otletae 132

- pauciflora 132

- petelotii 147

Hamularia 8, 43

Hancockia 7, 41

Hemipilia 6, 39

Herminium 6, 40

165

Herpysma 6, 38, 127, 130

- longicaulis 127

Hetaeria 6, 38, 127

- affinis 128, 131, 135, 138

- alta 128, 129, 137

- anomala 128, 130, 138

- biloba 128

- grandiflora 128

- hainanensis 128

- nitida 128, 131, 135, 138

- oblongifolia 128, 129, 135, 138

- parvifolia 155

- pauciflora 149

- poilanei 139

- rotundiloba 128

- rubens 131

- tokioi 149

Hippeophyllum 7, 42

Holcoglossum 8, 48

Hygrochilus 8, 46

Hysteria veratrifolia 93

Kingidium 8, 47

Lecanorchidinae 7

Lecanorchis 7, 15, 50

Limodoraceae 37

Liparis 7, 42

- nervosa 14

Listera 6, 38

Ludisia 6, 39, 132

- dawsoniana 132

- discolor 132, 135, 161

- discolor 147

- otletae 132

Luisia 8, 48

Macodes 6, 38, 133

- cupida 133, 135

- petola 133, 134

Malaxideae 7

Malaxidinae 7

Malaxis 7, 21, 42

Malleola 8, 49

Micropera 8, 46

Microsaccus 8, 45

Mischobulbon 7, 41

Monochilus flavus 157

- nervosum 154

Monomeria 8, 43


166

Myrmechis 6, 39, 134

- gracilis 134

- pumila 134, 135

Neogyna 7, 42

Neottia amoena 160

- australis 160

- petola 133

- procera 120

- sinensis 160

- viridiflora 124

Neottiaceae 37

Neottieae 6

Nephelaphyllum 7, 41

Nervilia 7, 40

Nervilieae 7

Neuwiedia 6, 37, 50

- annamensis 51

- balansae 51

- griffithii 52

- inae 52

- javanica 51

- veratrifolia 50

- zollingeri 50

- - var. annamensis 51, 163

- - var. balansae 51, 54, 55, 163

- - var. javanica 51, 55

- - var. singapureana 52

Oberonia 7, 42

- ensiformis 14

Oberoniinae 7

Odontochilus 63, 39, 50, 136

- acalcaratus 139, 141, 142

- brevistylus 139, 141, 145

- elwesii 136, 140, 142, 161

- flavescens 138

- inabai 139, 145, 146, 161

- lanceolatus 139, 143, 145

- poilanei 136, 139, 142

- pumilus 134

- saprophyticus 136, 140, 142, 161

- tortus 139, 144, 145, 146

- umbrosus 139, 143, 145, 161

Ophrys spiralis 158

Orchidaceae 6, 37

Orchideae 6, 37

Orchidinae 6, 15

Orchidoideae 6, 12, 22

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

Orchipedum 6, 38, 146

- echinatum 146, 150, 161

- plantaginifolium 146

Orchis 37

- strateumаtica 153

Ornithochilus 8, 45

Otochilus 7, 42

Pachystoma 7, 41

Panisea 7, 41

Paphiopedilum 6, 34, 37, 58

- aestivum 83

- × affine 86

- amabile 82

- angustatum 65

- appletonianum 61, 80, 81, 90

- × areeanum 72

- - × P. gratrixianum 86

- - × P. villosum 86, 87, 91

- armeniacum 62

- × aspersum 72

- barbatum 83

- barbigerum 60,70

- - var. aspersum 72, 89, 163

- - var. coccineum 71, 72, 73, 89

- - var. lockianum 72

- - × P. gratrixianum 86, 91

- callosum 61, 80, 83

- - var. angustipetalum 80

- - var. callosum 82, 85, 90

- - var. potentianum 82, 83, 90

- - var. sublaeve 82

- - var. warnerianum 82, 90, 163

- - × P. gratrixianum 86, 91

- - × P. villosum 86, 87, 91

- caobangense 73

- cerveranum 80

- chaoi 74

- chiwuanum 70

- coccineum 72

- concolor 14, 59, 67, 71, 88

- × cribbii 86, 87

- × dalatense 86, 87

- delenatii 59, 61, 64, 88

- delicatum 73

- dianthum 60, 84, 85, 91

- dollii 74

- emersonii 60, 67, 68, 88

- - inval. var. huonglanae 67

- esquirolei 69

- × fanaticum 92

- gratrixianum 61, 76

- - var. daoense 76, 77, 81, 89

- - var. gratrixianum 76, 89

- - × P. villosum 86, 91

- hainanense 80

- hangianum 60, 66, 68, 88

- helenae 60, 73, 75, 89

- - f. aureum 73

- - × P. hirsutissimum 86, 87, 91

- henryanum 61, 73, 74, 75, 89

- × herrmannii 86, 87

- hiepii 65

- hilmari 62

- hirsutissimum 60, 69

- - var. chiwuanum 69, 70, 71, 89

- - var. esquirolei 69, 71, 89

- huonglanae 67

- insigne 58, 73

- - var. barbigerum 72

- jackii 63

- - var. hiepii 65

- macranrhum 80

- malipoense 59, 62

- - var. hiepii 63, 64, 65, 88

- - var. jackii 63, 88

- - var. malipoense 63, 64, 88

- - × P. micranthum 91, 92

- micranthum 59, 65, 68, 88

- mirabile 62

- parishii var. dianthum 84

- potentianum 83

- purpuratum 61, 83, 85, 91

- purpuratum 86

- - var. hainanense 83

- rhizomatozum 72

- saccopetalum 69

- singchii 66, 67

- sinicum 83

- tranlienianum 60, 73, 75, 89

- tridentatum 80

- vejvarutianum 72

- vietnamense 59, 62, 64, 88

- villosum 61, 77

- - var. annamense 77, 78, 81, 90


Turczaninowia 2008, 11(1) : 5–168

- - var. boxallii 77, 79, 90

- - var. fusco-roseum 78, 79, 90

- - var. fusco-viride 78, 79, 90

- - var. gratrixianum 76

- - var. villosum 77, 78, 90

Papilionanthe 8, 47

Parapterocerаs 8, 49

Pecteilis 6, 40

Pelatantheria 8, 46

Pennilabium 8, 49

Peristylus 6, 40

Phaius 7, 41

Phalaenopsis 8, 48

Pholidota 7, 42

- rubra 14

Phreatia 8, 43

Physurus 117

- blumei 117, 118

- chinensis 118

- herpysmoides 117

Platanthera 6, 40

Pleione 7, 41

Plocoglottis 7, 41

Podochileae 7

Podochilinae 7

Podochilus 7, 42

Polystachya 7, 44

Polystachyinae 7

Porpax 7, 44

Pomatocalpa 8, 46

Pristiglottis 138

- saprophytica 140

- torta 144

- umbrosa 143

Pteroceras 8, 48

Pterygodium sulcatum 152

Renanthera 8, 46

Rhynchogyna 8, 47

Rhynchostylis 8, 47

Rhomboda 6, 39, 147

- lanceolata 147, 149, 156

- longifolia 147

- pauciflora 149

- petelotii 147, 150, 161

- tokioi 147, 149, 150, 162

Robiquetia 8, 47

Saccolabiopsis 8, 47

Sarcoglyphis 8, 46

Satyrium repens 118

Schoenomorphus 93

- capitatus 94

Schoenorchis 8, 46

Smitinandia 8, 46

Spathoglottis 7, 41

Spiranthes 6, 38, 92, 158

- amoena 160

- australis 160

- sinensis 159, 160, 162

- spiralis 158

- spiralis 160

Spiranthinae 6, 158

Spiranthoideae 5, 6, 8, 20, 92, 163

Staurochilus 8, 46

Stereochilus 8, 46

Stereosandra 7, 50

Sunipia 8, 43

Sunipiinae 41

Taeniophyllum 8, 45

Tainia 7, 41

- viridifusca 14

Thecopus 8, 44

Thecostele 8, 44

Thecostelinae 8

Thelasiinae 8

Thelasis 8, 43

Thrichoglottis 8, 46

Thrixspermum 8, 47

Thunia 7, 40

Thuniinae 7

Trias 8, 42

Trichotosia 7, 43

Tropidia 6, 38, 92, 93

- angulosa 93, 94, 98, 115

- curculigoides 27, 93, 94, 98, 115

- graminea 94

- pedunculata 94

Tropidieae 6, 92

Tuberolabium 8, 49

Uncifera 8, 47

Vanda 8, 47

Vandeae 8

Vandoideae 8, 12, 26

Vandopsis 8, 45

Vanilla 7, 40

167

Vanillaceae 37

Vanilleae 6

Vanallinae 7

Vietorchis 6, 50

Vridagzenia 151

Vrydagzynea 6, 38, 151

- albida 151, 152

- albida 151

- formosana 151, 156, 162

Zeuxine 6, 39, 152

- affinis 154

- bidupensis 153

- biloba 128

- bonii 153

- evrardii 153

- flava 153, 157, 159, 162

- goodyeroides 152, 153, 156, 162

- grandis 152, 154, 156, 162

- langbianensis 134, 138

- nervosa 153, 154, 156, 162

- parvifolia 153, 155, 156, 162

- strateumatica 152, 153, 156

- sulcata 152

- thanmoiensis 112

- tonkinensis 155

- vietnamica 157, 158

Zeuxinella 6, 39, 157

- vietnamica 157, 158, 159, 162

Yoania 7, 50


168

Averyanov L. The orchids of Vietnam illustrated survey. Part 1.

CONTENTS

PREFACE ........................................................................................................ 5

ILLUSTRATED GLOSSARY OF THE ORCHID

IDENTIFICATION TERMINOLOGY ................................................... 10

ACKNOWLEDGEMENTS ............................................................................... 35

KEY FOR IDENTIFICATION OF ORCHID GENERA

IN THE FLORA OF VIETNAM .......................................................... 37

SUBFAMILY APOSTASIOIDEAE ................................................................... 50

SUBFAMILY CYPRIPEDIOIDEAE ................................................................. 58

SUBFAMILY SPIRANTHOIDEAE .................................................................. 92

LITERATURE .................................................................................................. 163

INDEX OF LATIN NAMES ............................................................................ 164

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